Chapter 8
Barley Breeding History, Progress,
Objectives, and Technology
EUROPE
Wolfgang Friedt
1
POsiTiOn Of BARLEY PRODUCTiOn
AnD BREEDinG
Barley (Hordeum vulgare) is one of the leading
cereal crops of the world and it is clearly number
2 in Europe, next to bread wheat (Triticum aestivum). For example, in Germany, approximately
1.9 million hectares of spring and winter barley
contrast to 3 million hectares of winter wheat.
The remaining small grain cereals, that is, oats,
rye, and triticale, together cover around 1 million
hectares.
On a global scale, the barley area harvested
worldwide has declined from a maximum of more
than 80 million hectares in the 1970s to nowadays
less than 60 million hectares (Table 8.1.1). During
the same time, the cultivation area in Europe went
down from approximately 53 to 29 million hectares in 2008 (Table 8.1.2). This represents 51%
of the world barley hectarage (56.8 million hectares). The major European barley-growing countries, France, Germany, Russia, Spain, Ukraine,
and the United Kingdom, account for about
three-quarters of the total annual barley grain
production of about 105 million tons, representing 67% of the world production (157,644,721 t
in 2008). Almost two-thirds (62%) of the total
Barley: Production, Improvement, and Uses. Edited by Steven
E. Ullrich © 2011 Blackwell Publishing Ltd.
F
European barley production comes from the
European Union (EU) member states (EU 27)
covering 50% of the cultivation area in Europe.
The average grain yield per unit area in the EU
(27) is 25% higher than the European grand mean
(45.4 vs. 36.2 dt/ha; cf. FAO 2009). These igures
demonstrate the outstanding importance of barley
for Europe and of the European crop for global
barley grain production.
The high yield potential of modern cultivars
and improved cultivation practices basically allow
continuously increasing barley production in all
areas. For example, in Germany, the average
grain yield of spring barley on a farm scale has
grown from less than 2.0 t/ha in the early nineteenth century to a current level of 6.1 t/ha. Over
the last four decades, barley yield in Europe has
been increased by 60%, in Western Europe by
80% even. The corresponding relative yield
increases worldwide amount to 53% (Table 8.1.1;
FAO 2009).
In recent decades, the productivity of barley
cultivars has risen by an annual rate of approximately 1%–2%, which is due to (i) genetic breeding progress in terms of more productive
cultivars, (ii) more eficient disease and insect
control, (iii) improved fertilization schemes, and
(iv) improved agricultural production technology
(harvest, storage, etc.) in general. Yield increases
go along with a substantial improvement of
malting quality and signiicant enhancement of
agronomic traits leading to better yield stability,
for example, due to resistances against lodging,
diseases, and insects. Barley cultivars released
and grown in Europe today are characterized
by a considerable spectrum of resistance against
158
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Barley Breeding History, Progress, Objectives, and Technology
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Table 8.1.1 Barley production: comparison of global production igures with those of Europe (total and western), 1966–2008
(FAO2009)
World
Europe (Total)
Western Europe
Perioda
Area Harvested
(M ha)b
Grain Yield
(t/ha)c
Area Harvested
(M ha)b
Grain Yield
(t/ha)c
Area Harvested
(M ha)b
Grain Yield
(t/ha)c
1966–1968
1976–1978
1986–1988
1996–1998
2006–2008
Change (%)d
61.6
81.1
77.6
62.0
56.2
91
1.7
2.1
2.2
2.4
2.6
153
34.3
53.4
48.7
30.9
28.7
84
2.0
2.3
2.4
2.9
3.2
160
5.1
6.3
5.3
4.3
4.0
78
3.3
3.8
4.9
5.9
5.9
179
a
Averages of three consecutive years each.
106 ha.
c
Metric tons per hectare.
d
Relative values 2006–2008 versus 1966–1968.
b
virus diseases—particularly soil-borne yellow
mosaic viruses—and fungal pathogens such
as powdery mildew (Blumeria graminis), leaf
rust (Puccinia hordei), or scald (Rhynchosporium
secalis). The number of resistant cultivars has
been signiicantly increased throughout the past
few decades (Table 8.1.3).
For example, a comparison of rankings for
powdery mildew attack and grain yield of current
winter barley cultivars with those on the market
in Germany two or three decades ago clearly
shows the drastic change in the varietal performance: some modern winter barley cultivars even
combine a very high yield potential with a pronounced mildew resistance, which was not achievable before (cf. Friedt et al. 2000; Anonymous
2009). The main reason for strongly improved
mildew resistance particularly in the spring cultivars is seen in the wide use of effective resistance
genes. Various genes have been exploited throughout the decades with some emphasis on the Mla
locus complex. Recently, however, the interest of
breeders has focused on the mlo gene. Today,
probably more than one third of the spring barley
cultivars in Europe carry this gene conferring
practically complete ield resistance against
powdery mildew.
A similar situation also exists with regard to
resistances against other diseases, such as scald
(R. secalis), leaf or brown rust (P. hordei), and
yellow mosaic-inducing viruses (barley mild
mosaic virus [BaMMV] and BaYMV), some of
the most important diseases of winter barley in
Europe. Whereas in the 1980s only a few cultivars
showed satisfactory resistance, many cultivars
expressing a high level of resistance, that is, low
susceptibility (1–3 on a 1–9 scale according to the
German Plant Variety Ofice) have been released
more recently (Table 8.1.3). For improving rust
and scald resistance, various genes have been
used. For example, Rph7 provides fairly stable
leaf rust resistance and has been transferred
to various cultivars. New genetic sources for
scald resistance have been identiied showing
low disease incidence in different regions of
Europe (cf. Friedt and Rasmussen 2004). Also,
Pyrenophora net blotch and Ramularia leaf blotch
have to be mentioned as spreading fungal diseases, which occasionally and regionally cause
high infections and damage to European barley
crop.
The increased level of disease resistance in
present-day cultivars also relects a general tendency in agricultural practice, as farmers are
under a constantly increasing pressure to reduce
the use of agrochemicals and pesticides for environmental reasons, thus enforcing focus on the
resistance properties of varieties. Therefore,
breeding for resistance in barley is a very important task on a global scale, since average yield
losses worldwide of up to 30% have to be faced
due to fungal and viral diseases along with insect
2
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Barley: Production, Improvement, and Uses
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Table 8.1.2
Major igures of barley production in European countries in 2008 (FAO2009)
Country
131
Albania
Austria
Belarus
Belgium
Bosnia and Herzegovina
Bulgaria
Croatia
Czech Republic
Denmark
Estonia
Finland
France
Germany
Greece
Hungary
Ireland
Italy
Latvia
Lithuania
Luxembourg
Malta
Moldova
Montenegro
The Netherlands
Norway
Poland
Portugal
Romania
Russian Federation
Serbia
Slovakia
Slovenia
Spain
Sweden
Switzerland
Macedonia a
Ukraine
United Kingdom
Total or average
European Union (EU) (EU 27)
132
Area Harvested (ha)
1400
185,857
612,639
55,016
22,723
222,659
61,200
482,395
717,300
141,200
585,500
1,799,300
1,961,700
150,000
332,000
181,200
330,067
131,200
332,500
9739
400F
130,179
790
50,200
128,240
1,206,560
43,100
386,706
9,420,800
92,417
213,050
19,229
3,462,400
407,700
33,112
47,351
4,167,200
1,032,000
29,157,029
14,473,752
Production (t)
25.0
52.1
36.1
77.0
33.4
39.4
44.5
46.5
47.3
24.7
36.4
67.6
61.0
25.3
44.5
69.0
37.5
23.4
29.2
54.2
40.0
27.1
15.2
61.8
41.3
30.0
23.2
31.3
24.6
37.3
41.8
39.9
32.5
44.2
61.2
34.4
30.3
59.5
3500
967,921
2,212,480
423,800
75,841
878,000
272,100
2,243,865
3,396,000
349,100
2,128,600
12,171,300
11,967,100
380,000
1,478,200
1,249,700
1,236,697
307,100
970,400
52,816
1600
353,124
1200
310,200
530,000
3,619,460
99,800
1,209,410
23,148,450
344,141
891,317
76,788
11,261,100
1,801,000
202,700
162,779
12,611,500
6,144,000
36.2
45.4
105,533,089
65,662,080
Source: http://faostat.fao.org/.
a
Former Yugoslav Republic of Macedonia.
Major seed producers (1000 t): Belarus (250), Czech Republic (104), Denmark (132), Finland (136), France (235), Germany
(322), Poland (235), Russia (3000), Spain (850), Ukraine (700), and the United Kingdom (143). The global annual seed
production varied from 9.5 to 9.8 million tons in 2006–2008. Of the total global annual seed production, 6.8 (70%) is
generated in Europe and 2.8 MT (29%) in the EU (27).
pests. Therefore, in order to avoid necessary
applications of chemicals for the prevention of
respective yield losses and to ensure economic
grain production, future barley breeding for
disease and pest resistances should even be further
intensiied.
F
Grain Yield (dt/ha)
CURREnT sPRinG BARLEY CULTiVARs
COMBininG HiGH QUALiTY AnD
DisEAsE REsisTAnCE
Modern European spring barley varieties trace
back to European landraces from Bavaria (south-
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Barley Breeding History, Progress, Objectives, and Technology
Table 8.1.3
diseasesa
161
Number of winter barley cultivars listed in Germany expressing pronounced resistances to major pathogens and
Year
No. of Listed Cultivars
Mildew Resistanta
Scald Resistanta
Brown Rust Resistanta
Yellow Mosaic Resistanta
1980
1986
1994
2002
2009
35
46
69
84
67b
3
3
14
32
33
3
1
5
28
18
0
0
3
18
19
n.d.
n.d.
19
49c
52c
Source: Beschreibende Sortenliste, Bundessortenamt, Hannover, Germany (German Plant Variety Ofice).
a
Low susceptibility: scores 1–3 (scale 1–9: 1 = minimum susceptibility, 9 = maximum susceptibility) for powdery mildew
(B. graminis), scald (R. secalis), brown rust (P. hordei), and yellow mosaic viruses (BaYMV and BaMMV).
b
Comprising 34 six-row and 33 two-row cultivars.
c
Four cultivars each are also resistant against BaYMV-2.
n.d., not detected.
ern Germany), Moravia (today Czech Republic),
Sweden, and the United Kingdom. Cycles of
cross-breeding irst focused on hybridizations
among the landraces, leading, for example, to
early varieties like “Isaria” (1924) and “Kenia”
(1931). Later, more distant and exotic barley
materials were included, for example, Hordeum
laevigatum and “arabische,” two major sources of
disease resistance inally leading to outstanding
spring varieties like “Aramir” (released in
Germany 1974) and the mlo11-carrying variety
“Apex” (1983). A third cycle of cross-breeding
comprised the Czechoslovakian short-straw,
heavy-tillering mutant “Diamant” (Fig. 8.1.1),
which also contributed to better malt quality due
to high proteolytic enzyme activity (Fischbeck
3 1992, cit. Friedt et al. 2000). This led to the
important cv. “Trumpf” (1973), also named
“Triumph,” which acquired a wide distribution
in Europe. Numerous progenies of crosses
between Trumpf and representatives of the
“Aramir group” became popular cultivars. Major
members of this cycle were “Blenheim” (UK,
1985), “Carmen” (A, 1985), “Prisma” (NL, 1985),
“Natasha” (F, 1986), and “Cheri” (D, 1987). An
outstanding exponent of this variety group is
“Alexis” (D, 1986), a widely grown cultivar with
additional genes for mildew resistance, for
example, the mlo gene of the Diamant mutant
“Helena” (Fig. 8.1.1). On the other hand, the
improved quality of Alexis as compared to
Trumpf was probably derived from parents like
133
“Proctor” or Isaria (Fischbeck 1992, cit. Friedt
et al. 2000). More recent outstanding spring
barley cultivars were “Scarlett” (1995) and
“Barke” (1996), both classiied at maximum malt
quality (score 9) and thereby representing an
improvement in comparison to Alexis (score 8).
“Marnie,” a later spring malt barley cultivar,
combines very high grain yield (ca. 10% superior
to Barke) with improved malting quality (superior
lytic parameters) and resistances against powdery
mildew, leaf rust, scald, and net blotch. Marnie’s
outstanding disease resistance derives from Israeli
H. spontaneum used as a cross parent (J. Breun,
pers. comm.).
Malting quality is an economically very important but also genetically complex trait, comprising
a number of different quality parameters. For a
reliable description of this complex and its stability, multilocation trials are necessary, which need
to be conirmed over several years of cultivation.
As many as 29 out of 47 (62%) spring barley
cultivars in Germany are classiied by very high
malting quality (extract content) represented by a
score of 8–9 (Anonymous 2009). Malt quality is
deined by the following speciic criteria (desired
expression): Kohlbach index, that is, enzyme
activity (high), extract difference (low), extract
content (high), and protein content (low). Results
of EBC trials with European varieties released
during the past few decades clearly show that
modern, high-yielding cultivars have a drastically
increased Kohlbach index and extract content
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Barley: Production, Improvement, and Uses
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Isaria (24)
Rika
(51)
Baladi
~~~~~
Ragusa
R
~~~~~~
(29)
Rika (51)
CP
Sulu
S l
~~~~
Diamant (65)
****
RiBaRi
Helena (83)
Proctor
(52))
(5
St
****
F1 1400
W409 /250
U1 1455/12714
Drossel/RiBaRi
Him T253/Volla
Him.T253/Volla
Br. 1453e16
Aufis (74)
Trumpf (73)
Br. 1622d54
Br . 2629
(Piruette/
Br. 1622)
Br. 2476
Br. 1400
W1455
(Helena/RiBaRi) (W409/Proctor)
Georgie (77)
Br. 1747
Rupee
Br. 1622
W5604
(Medusa/
Diamant)
Hanna
Alexis (86)
Br. 2730
Kym (81)
Libelle (90)
Amazone
(86)
Br. 3791
Scarlett (95)
Barke (96)
_____
~~~~
*****
( ) Year of release
Selection from landraces
Source of resistance
Mutation
fig. 8.1.1. Pedigree of major German spring barley cultivars as important parents of modern high malting quality
varieties.
along with a strongly reduced extract difference
and a lower protein content (R.S. Schildbach,
pers. comm.). These are the reasons for the
observed progress regarding extract content as
the major overall quality trait.
However, the barley grain is not only used for
feeding and beer production but also for producing malt whisky. Breeding lines have been developed, which combine all the relevant trait
expressions desired for malt whisky production,
for example, sprit yield, enzymatic potential
(diastatic power), and low content of glycosidic
nitrile. Cultivar “Optic,” derived from the cross
Chad × (Corniche × Force), has been the leading
spring barley in England and in Scotland. Its successors, “Publican” and “Forensic,” are considered to be particularly suitable for British
(Scottish) conditions, helping growers with extra
yield and also providing quality advantages to end
F
users, for example, distillers (Barley Malt News
5
2009; http://www.newfarmcrops.co.uk).
Modern variety breeding aims at the development of cultivars with high adaptability and elasticity so that they may be successfully grown in
widely different environments. Therefore, cultivars developed under continental climatic conditions as prevalent in large parts of Germany
should also be competitive in other European
countries and elsewhere. Such successful modern
cultivars represent optimal genetic complexes
determining the necessary trait combinations
providing high crop stability and performance. As
shown in Table 8.1.4, actual German spring
barley cultivars are characterized by such optimal
trait compositions, for example, “Streif” or
“Tocada,” combining broad resistances against
abiotic and biotic stresses with high yield poten6
tial and superior malting quality.
4
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Table 8.1.4 Major positive traits of spring barley cultivars listed in Germany in 2009
Cultivar (Year of Release)
Disease Resistance
Crop Stability
Grain Yield
Malt Extract
Annabell (1999)
Braemar (2002)
Average
Average
High
High–very high
High-very high
High
Average-high
Very high
High
High–very high
Average-high
Very high
Average-high
High–very high
Grace (2008)
Marthe (2005)
Quench (2006)
Streif (2007)
Tocada (2008)
Moderate BR and NB resistance
Resistant: PM (high), BR
(moderate)
Broad partial resistances
Broad resistance, particularly to
PM and NB
Resistant: PM (high), scald
(moderate)
Broad resistance, particularly
against PM and NB
Moderate NB and BR resistance
Low culm and ear
cracking
Low lodging, ear and
culm cracking
Low lodging, ear and
culm cracking
Low lodging, ear and
culm cracking
Low lodging, ear and
culm cracking
Low lodging, ear and
culm cracking
134
BR, brown rust (P. hordei); NB, net blotch (Pyrenophora teres); PM, powdery mildew (B. graminis).
CURREnT WinTER BARLEY VARiETiEs
COMBininG HiGH YiELD AnD
MALTinG QUALiTY
Malta
Carsten 1565 Strengs Aurea(SG)
W1106
Br. 127a14
Br. 59c21
Alpha
Dea
Herfordia
SvP 674
Malta
Modern European winter barley varieties are
mainly traced back to two sources of six-row
barley landraces, one from The Netherlands
and one from the Canadian winter barley sort
“Mammut.” The variety “Friedrichswerther
Berg” (1904), a progeny of hybridization between
these two pools, occupies a central position
in Central European winter barley breeding.
Subsequent breeding cycles based on combinations of six-row winter types and both twoand six-row spring barleys, including, for
example, the mildew-resistant Dalmatian landrace “Ragusa,” led to a series of important varieties such as “Dea” (1953) and “Dura” (1961).
Much later, it became obvious that Ragusa also
imparted resistance against BaMMV to the
German winter barley gene pool. “Vogelsanger
Gold,” another major variety derived from a cross
of a mildew-resistant spring barley and repeated
backcrosses, together with cv. Dura, were major
cross parents for later widely grown six-row
winter barleys like “Corona” (1980) and more
recent cultivars (cf. Fischbeck 1992, cit. Friedt
et al. 2000). Varieties like Dea were also parents
of crosses with two-row spring barley cultivars
like “Ingrid,” which inally in 1968 led to the
development of cv. “Malta” (see Fig. 8.1.2), the
Br. 301a
W5907
Br. 269c22
Marinka (85)
Labea (92)
Angora (90)
Tiffany (96)
Jura (95)
Regina (95)
fig. 8.1.2. Pedigree section of important two-rowed
German winter barley cultivars as major parents of modern,
quality-improved two-row winter barley varieties (J. Breun,
pers.comm.).
major parent of subsequent widely distributed
two-row winter-type malting barley cultivars
such as “Sonja” (1974), “Igri” (1976), “Marinka”
(1985), “Trixi” (1987), and progenies thereof.
A more recent cross between the two-row
winter barleys “Labea” (1992) and Marinka led
to cultivars like “Jura” (1995) and “Tiffany”
(1996) (Fig. 8.1.2; cf. Friedt et al. 2000).
Particularly the latter two closely related cultivars
rapidly gained importance in Germany in the late
1990s, since they combined a fairly good malt
quality (extract content 7, equivalent to spring
barley Apex) with a suficient yield performance
(cf. Friedt and Rasmussen 2004). Winter malting
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Barley: Production, Improvement, and Uses
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barley cultivation in Western Europe is still
expanding at the expense of spring barley. Among
the currently leading commercial winter two-row
varieties are “Campanile,” “Finesse,” “Finita,”
“Malwinta,” “Passion,” and “Reni.” Examples of
currently leading six-row winter types are
“Fridericus,” “Highlight,” “Lomerit,” and
“Naomie.” It is worth mentioning that hybrid
breeding is gaining importance in European
winter barley breeding. For example, the highyielding six-row hybrid cv. “Zzoom” was added
to the German variety list in 2008 (Anonymous
2009).
BROAD-BAsED ViRUs-REsisTAnT
WinTER BARLEY CULTiVARs
Considerable breeding progress has been made
with regard to virus resistance of European barley
varieties. Besides barley yellow dwarf virus
(BYDV), particularly yellow mosaic-inducing
soil-borne viruses transmitted by the plasmodiophorid Polymyxa graminis have recently expanded
due to the extension of cereal cultivation and
infested acreage. They cause damage to barley
crops in many European growing regions. Among
the insect-transmitted viruses vectored by aphids,
BYDV has an outstanding relevance for barley
cultivation and is expected to gain importance
with advancing global warming. There is no
doubt that breeding of virus-resistant cultivars is
the best, cost-effective, and environmentally
compatible approach to prevent virus infection.
Numerous sources of resistance or tolerance to
the major barley viruses were identiied in the
past few decades and have been used in classical
breeding as well as by molecular breeding
approaches (cf. Ordon et al. 2009).
Since these viruses were identiied in the late
1970s, the breeding goal of high-yielding virusresistant cultivars has been achieved quite rapidly
(cf. Werner et al. 2000), starting from the identiication of resistant sources within exotic cultivars
or unadapted germplasm and followed by usually
biparental crosses to combine resistance with
other agronomic and quality traits. At the beginning of the 1990s, some important cultivars like
F
“Jana” were already resistant to BaMMV and
BaYMV. Furthermore, extensive screening and
genetic analysis enabled the release of cv. “Tokyo”
([(Fallon × 13060) × 87 – 5381 B] × Swift]; R.
Hemker, pers. comm.), only a few years after the
irst detection of the resistance-breaking strain
BaYMV-2 in Europe. The resistance of “Tokyo”
was derived from line number 13060, which has
“Mokusekko 3,” the donor of rym5 in its pedigree. Due to its agronomic shortcomings, susceptibility to scald, and the relatively limited
distribution of BaYMV-2, this cultivar has not
achieved great acceptance by growers. However,
resistant lines derived from Tokyo with better
agronomic performance are being released.
Winter barley is particularly attractive to
growers due to its high yield potential as compared to spring types. On the other hand, due to
the more recent history of this crop, the variation
for disease resistance or malting quality is rather
limited in comparison to spring barley. Nevertheless, great improvements have been achieved
as may be recognized from Table 8.1.5. Recently
listed German winter barley cultivars such as
“Jade” (two-row) and “Christelle” (six-row)
exhibit superior trait compositions, combining
quantitative resistances against major diseases
with outstanding yield and promising malting
quality. From that, it can be expected that future
varieties will combine their maximal yield potential with suficient stability (e.g., lodging and
stress resistances) as well as suficient malting
quality. Ultimately, such a trend can be expected
to entirely replace spring barley with winter
barley in many European barley-growing areas.
BiOTECHnOLOGY-BAsED AnD
MARKER-AssisTED APPROACHEs:
EVOLUTiOn Of BREEDinG METHODs
Acceleration of barley breeding
via haploidy
Combinations of different resistance genes or the
introgression of novel resistances from nonadapted germplasm into adapted cultivars’
background are classically achieved by sexual
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Table 8.1.5
165
Major positive traits of new winter barley cultivars, released in Germany in 2009
Cultivar
Disease Resistance
Crop Stability
Grain Yield
Grain Quality
Christelle (six-row)
Mildew, net blotch, brown rust,
BYMV
Mildew, brown rust, BYMV
Mildew, BYMV
Mildew, net blotch, scald,
brown rust, BYMV
Mildew
Low culm cracking
Very high
Moderate test weight
Low spike cracking
Very high
Very high
Very high
Low–moderate test weight
Moderate–high test weight
Low–moderate test weight
Low culm and spike
cracking
Lodging resistant
Lodging resistant,
low culm cracking
Low spike cracking
High-very high
Moderate test weight
High
High
High test weight
Moderate–high test weight
Moderate-high
Moderate–high test weight
Low culm cracking
High
Moderate test weight
Kathleen (six-row)
Semper (six-row)
Souleyka (six-row)
Anisette (two-row)
Canberra (two-row)
Jade (two-row)
Lucie (two-row)
Zephyr (two-row)
Mildew, BYMV
Mildew, net blotch, scald,
BYMV
Mildew, scald, brown rust,
BYMV
Scald, BYMV
BYMV, resistant against barley yellow mosaic viruses.
recombination, that is, crosses between selected
parental lines followed by phenotypic selection in
the segregating offspring. In this case, the success
of breeding entirely depends on extensive ield
and/or greenhouse tests for resistance to the
respective pathogen(s). However, since barley is
damaged by many pathogens, which often show a
rapid adaptation to their hosts’ resistance, breeding for resistance is a very complex task, and the
identiication of desired recombinants by phenotypic selection, for example, in pedigree selection
schemes, has almost reached the limits of manageability. Thus, methods of plant biotechnology
like anther and microspore culture allowing the
rapid production of homozygous doubled-haploid
(DH) lines and cultivars were highly welcome and
have been implemented into barley breeding
schemes. For example, via anther culture, the
spring barley cv. “Henni” (D, 1995), the two-row
winter barley “Anthere” (D, 1995), and the sixrow cultivars “Uschi” (D, 1997), “Sarah” (D,
1997), “Carola” (F, 1997) and “Nelly” (D, 1998)
had been released earlier than expected and
became widely grown in Europe (E. Laubach,
pers. comm.). It is obvious that a substantial time
gain can be achieved by the application of this
biotechnology step in barley breeding (Fig. 8.1.3).
Consequently, the “haploid breeding method” is
widely applied now and has gained great importance for barley breeding as demonstrated by the
fig. 8.1.3. Generalized breeding scheme for spring and
winter barley assisted by the application of the “haploid
method” (E. Laubach, pers. comm.).
127
growing number of DH varieties released (E.
Laubach, pers. comm.).
Molecular markers and
marker-assisted selection
In addition, the development of molecular
markers, which to some extent allow the transfer
of selection steps from the phenotypic (ield)
to the genotypic (laboratory) level, offers new
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F
opportunities for a more eficient barley breeding
aiming at desired combinations of resistance,
yield, and quality (Friedt et al. 2002).
For example, to enhance the resistance against
BYDV, Habekuss et al. (2009) used DH lines and
molecular markers to combine resistance genes
Ryd2 and Ryd3 and a quantitative trait locus
7 (QTL) from cv. “Post” on chromosome 2H. DH
lines combining Ryd2 and Ryd2 are reported to
show lower virus titer and reduced symptom
expression as compared with parental materials.
Another example is yellow mosaic-inducing
viruses: barley cv. Taihoku A has been described
as resistant to yellow mosaic viruses reported in
Germany (i.e., BaMMV, BaYMV, BaYMV-2,
and BaMMV-Teik). This cultivar carries the
BaMMV resistance gene rym13 on chromosome
4H, which may be responsible for the resistance
against the whole virus complex. By bulked segregant analysis using simple sequence repeats
(SSRs) and ampliied fragment length polymorphisms (AFLPs), Humbroich et al. (2009) constructed a map with the closest marker linked to
the BaMMV/BaMMV-Teik resistance locus at a
distance of 1 cM. These markers are useful tools
to introduce resistance to BaMMV-Teik into
adapted breeding lines carrying other resistance
genes, which are not effective against all BaMMV/
BaYMV strains known in Europe.
For a long time, powdery mildew has been
known as an important pathogen of barley in
almost every growing region so that resistance
against mildew is of utmost importance. To identify molecular markers, Korell et al. (2008) used
a cDNA-AFLP approach to study near-isogenic
barley lines differentiated by alleles of the resistance gene Mlg located on chromosome 4H.
Based on the identiication of a short differential
fragment (37 bp), which turned out to be part of
8 a nucleoside diphosphate kinase, a CAPS marker
cosegregating with Mlg was developed. Due to its
codominance, clear banding pattern, and close
linkage, this marker is well suited for markerassisted selection procedures.
While DNA marker analyses gain increasing
importance in plant breeding and become more
widely adopted in cultivar development, the
capacity for high-throughput analyses at low cost
is crucial for its practical application. Automation
of the analysis processes is a way to meet these
requirements. For this purpose, the company
Svalöf Weibull AB, Sweden, has developed a
fully automated PCR system. It was evaluated on 9
barley lines and was shown to be capable of analyzing up to 2200 samples per day at costs of EUR
0.24 per analysis for marker-assisted selection and
quality control of genetically modiied plants
(Dayteg et al. 2007).
10
Genome analysis and GM barley
Today, sequencing of the barley genome has
become a realistic option. Necessary steps to
establish and improve genomics tools have been
initiated and assembled, coordinated by an international consortium (cf. Schulte et al. 2009). A
suitable reference genome sequence will be an
excellent foundation for “selection with markers
and advanced recombination technology”
(SMART) or marker-assisted breeding, leading
to future genomics-based barley improvement.
In the foreseeable future, access to highly
sophisticated breeding tools with a broad genetic
diversity (cf. Fig. 8.1.4) as an absolute basis for
gain of selection will probably become the major
limiting factor for further breeding progress.
Therefore, the conservation, maintenance, and
evaluation of plant genetic resources are urgently
needed social tasks to build the foundation for
future reasonable plant breeding, crop improvement, and entirely successful agriculture. This
will be even more the case if expected climate
changes will lead to additional requirements of
new varieties such as drought or heat resistances.
Within the international network of gene banks,
for example, the Institute of Plant Genetics
and Crop Plant Research (IPK), Gatersleben,
Germany, plays a major role not only for barley
conservation but also for barley genomics (see
Stein et al. 2007; Schulte et al. 2009; cf. Sato
et al. 2009).
Genetic transformation is not only necessary
for developing new cultivars with speciically
modiied traits such as improved disease and
stress resistance or grain quality. Stable genetic
transformation represents also an optimal
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1
2
3
4
5
6
10 cm
167
10 cm
10 cm
10 cm
10 cm
10 cm
10 cm
10 cm
Barley Breeding History, Progress, Objectives, and Technology
7
8
fig. 8.1.4. Barley spike phenotypes: 1, two-rowed cv. “Barke” (wild type); 2–8, mutant phenotypes; 2, awnless; 3, intermedium erectoides; 5, brachytic; 6–8, “irregular” spikes. Source: Dr. S. Gottwald.
approach to the detailed elucidation of plant gene
function(s). This is considered particularly relevant in barley, a widely used experimental model
plant. So, Hensel et al. (2008) presented details of
the establishment of a protocol for Agrobacteriummediated gene transfer to immature embryos,
enabling the highly eficient generation of transgenic barley. Advancements were made by comparing the inluence of various treatments and
cocultivation conditions, leading to transgenic
lines expressing the respective transgene product
at low T-DNA copy numbers. In particular, the
newly established protocol turned out to be useful
for transformation of various spring and winter
barley genotypes besides the highly amenable cv.
“Golden Promise.” A very useful tool is now
available for functional gene analyses as well as
genetic engineering approaches in barley cultivar
development.
Nevertheless, it cannot be expected that transgenic barley will play a major role in European
agriculture in the near future. Strong reservations
against GM crops and GM food among some
consumers and farmers have caused political decisions by European authorities to the disadvantage
of the application of GM technology up to the
present.
128
COnCLUsiOns AnD PERsPECTiVEs
Barley breeding in Europe has been extremely
successful throughout the last century. Cycles of
cross-breeding, which irst made use of hybridization between European landraces, and later
exploiting more distant germplasms providing
valuable disease resistances, and inally combining high-yield and high-quality varieties, have led
to highly productive modern cultivars, both in
spring and winter barley. In most cases, the practical breeding techniques comprised little more
than manual hybridization, careful observation,
precise testing, and conscious selection. More
recently, breeding programs have been enhanced
by the implementation of modern biotechnology
tools, like the “haploid method.” Now, it is
becoming obvious that marker-assisted selection
procedures—particularly PCR-based techniques
combined with fast and high-throughput analysis,
also called SMART breeding—will further
enhance the process of selecting resistant varieties
with superior agronomic performance in the
future. As a consequence, existing breeding
schemes including “fast track” procedures will
even be accelerated and carried out more
eficiently than up to now.
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Barley: Production, Improvement, and Uses
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In addition to methodological progress, “new”
traits including special qualities (”functional
barley”) for alternative, for example, chemical
purposes, such as high amylose and waxy
barley, would potentially allow the diversiication
and extension of barley cultivation and use.
Furthermore, hulless barley may help to improve
feed quality regarding digestability and nutrient
contents, probably combined with phytase activity or low phytate content for better phosphate
use. In this regard, a better understanding of gene
and genome functions will enable more straightforward breeding approaches like the direct
transfer of donor genes into suitable receptor
cultivars.
In this context, it is worth mentioning that a
11 new EMS-induced mutant population comprising 10,279 M2 individuals of two-rowed spring
barley cv. Barke was described recently (Gottwald
12 et al. 2009). This new TILLING resource has
potential for use in fundamental research as well
as in applied breeding.
ACKnOWLEDGEMEnTs
Expertise on varieties and pedigrees provided by
Josef Breun, Herzogenaurach, Germany, and Dr.
Eberhard Laubach, Nordsaat, Germany, is gratefully acknowledged. I wish to thank Prof. Dr.
Frank Ordon for detailed information regarding
disease resistance and marker-assisted selection.
REfEREnCEs
F
Anonymous. 2009. ••. In •• Bundessortenamt (ed.).
Beschreibende Sortenliste 2009. Landbuch-Verlag,
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Hannover, Germany.
Behn, A., L. Hartl, G. Schweizer, G. Wenzel, and M.
Baumer. 2004. QTL mapping for resistance against nonparasitic leaf spots in a spring barley doubled haploid
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population. Theor. Appl. Genet. 108:1229–1235.
Dayteg, C., S. Tuvesson, A. Merker, A. Jahoor, and A.
Kolodinska-Brantestam. 2007. Automation of DNA
marker analysis for molecular breeding in crops: practical
experience of a plant breeding company. Plant Breed.
126:410–415.
Food and Agriculture Organization (FAO) of the United
Nations. 2009. Agricultural Statistics Web site, available at
http://faostat.fao.org/.
Friedt, W. and M. Rasmussen. 2004. ••. Symp. EUCARPIA
Cereal Section, Salsomaggiore, Italy, •• ••, ••.
Friedt, W., K. Werner, and F. Ordon. 2000. ••. pp. 271–279.
In •• •• (ed.). Proc. 8th Int. Barley Genet. Symp., Adelaide,
Vol. 1. ••, ••.
Friedt, W., K. Werner, B. Pellio, C. Weiskorn, M. Krämer,
and F. Ordon. 2002. Strategies of breeding for durable
disease resistance in cereals. Prog. Bot. 64:138–167.
Gottwald, S., P. Bauer, T. Komatsuda, U. Lundqvist, and
N. Stein. 2009. TILLING in the two-rowed barley cultivar “Barke” reveals preferred sites of functional diversity
in the gene HvHox1. BMC Res. Notes 2:258. doi:
10.1186/1756-0500-2-258.
Habekuss, A., C. Riedel, E. Schliephake, and F. Ordon. 2009.
Breeding for resistance to insect-transmitted viruses in
barley—an emerging challenge due to global warming.
J. f. Kulturplanzen 61:53–61.
Hensel, G., V. Valkov, J. Middlefell-Williams, and J.
Kumlehn. 2008. Eficient generation of transgenic barley:
the way forward to modulate plant–microbe interactions.
J. Plant Physiol. 165:71–82.
Humbroich, K., H. Jaiser, A. Schiemann, P. Devaux, A.
Jacobi, L. Cselenyi, A. Habekuss, W. Friedt, and F.
Ordon. 2009. Mapping of resistance against barley mild
mosaic virus-Teik (BaMMV)—an rym5 resistance
breaking strain of BaMMV—in the Taiwanese barley
(Hordeum vulgare) cultivar Taihoku A. Plant Breed. doi:
10.1111/j.1439-0523.2009.01721.x.
Korell, M., T.W. Eschholz, C. Eckey, D. Biedenkopf, K.H.
Kogel, W. Friedt, and F. Ordon. 2008. Development of a
cDNA-AFLP derived CAPS marker co-segregating with
the powdery mildew resistance gene Mlg in barley. Plant
Breed. 127:102–104.
Ordon, F., U. Kastirr, F. Rabenstein, and T. Kühne. 2009.
Virus resistance in cereals: sources of resistance, genetics
and breeding. J. Phytopathol. 157:535–545.
Sato, K., N. Nankaku, and K. Takeda. 2009. A high-density
transcript linkage map of barley derived from a single
population. Heredity 103:110–117.
Schmalenbach, I., J. Léon, and K. Pillen. 2009. Identiication
and veriication of QTLs for agronomic traits using
wild barley introgression lines. Theor. Appl. Genet. 118:
483–497.
Schulte, D., T.J. Close, A. Graner, P. Langridge, T.
Matsumoto, G. Muehlbauer, K. Sato, A.H. Schulman, R.
Waugh, R.P. Wise, and N. Stein. 2009. The International
Barley Sequencing Consortium—at the threshold of
eficient access to the barley genome. Plant Physiol. 149:
142–147.
Stein, N., M. Prasad, U. Scholz, T. Thiel, H. Zhang, M.
Wolf, R. Kota, K.R. Varshney, D. Perovic, I. Grosse, and
A. Graner. 2007. A 1000-loci transcript map of the barley
genome: new anchoring points for integrative grass
genomics. Theor. Appl. Genet. 114:823–839.
Werner, K., B. Pellio, F. Ordon, and W. Friedt. 2000. ••.
Plant Breed. 119:517–519.
Werner, K., W. Friedt, and F. Ordon. 2005. Strategies for
pyramiding resistance genes against the barley yellow
mosaic virus complex (BaMMV, BaYMV, BaYMV-2).
Mol. Breed. 16:45–55.
15
16
17
18
19
20
21
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Werner, K., W. Friedt, and F. Ordon. 2007. Localisation and
combination of resistance genes against soil-borne viruses
of barley (BaMMV, BaYMV) using doubled haploids and
22
molecular markers. Euphytica 158:323–329.
NORTH AMERICA
Rich D. Horsley and Bryan L. Harvey
Large breeding programs for barley in North
America exist in Canada, Mexico, and the United
States. This portion of the chapter will focus on
the programs in Canada and the United States
because of the working knowledge and associations the authors have with these programs and
the individuals working on them. The two largest
programs in Mexico are the national program,
which is the Instituto Nacional de Investigaciones
Forestales, Agricolas, y Pecurias (INIFAP), and
the program overseen by the International Center
for Agricultural Research in the Dry Areas
(ICARDA). For many years, the ICARDA
program was located at the headquarters of the
International Maize and Wheat Improvement
Center (CIMMYT) in El Batan, Mexico. The
national program has been successful in developing malting barley cultivars for Mexico, such as
Esmerelda, and the ICARDA program has been
successful in developing germplasm with multiple disease resistance for use around the world. In
2007, the ICARDA barley breeder responsible for
Mexico and South America was moved to the
ICARDA headquarters in Aleppo, Syria; however,
the responsibilities of this breeder for the region
did not change.
169
BARLEY PRODUCTiOn in CAnADA
Barley in Canada was introduced to New France
by the irst governor, Samuel de Champlain, in
1606. Canada’s irst brewery was built in Quebec
city in 1668 (Metcalfe 1995). Later, the British
brought two-row brewing barleys to Upper
Canada. Trade between Canada and the United
States thrived and brought good returns to
Ontario growers for their two-row malting barley.
This trade was brought to an end by U.S. protectionist measures in the form of the McKinley
Tariff Act in 1890. Therefore, Ontario growers
were forced to turn to feed markets for their
barley. Although the two-row barley cultivars
were highly suited to malting, they were low
yielding, and thus improvements were required
to facilitate viable barley production in that area.
The Hudson Bay Company introduced cereal
grains, including barley, to their trading posts
throughout Northwest Canada in the 1600s in an
attempt to attain food self-suficiency at these
posts. However, signiicant commercial production did not begin in Western Canada until the
arrival of Selkirk settlers in southern Manitoba in
1812. Barley has been a major cereal grain in
Western Canada since that time.
Over 90% of Canada’s barley acreage is in
Western Canada (Table 8.2.1). However, it is still
an important crop in Eastern Canada where
growing conditions are highly variable with a
multiplicity of agro-ecoregimes.
Canada today can be divided into speciic
barley-growing regions (Fig. 8.2.1). Below are
Table 8.2.1 Canadian barley production in thousands of tons, 2004–2008 (calculated from data provided by Statistics
Canada; http://www.statcan.gc.ca; veriied July 22, 2009)
Year
Area
2004
2005
2006
2007
2008
Mean
%
Canada
Maritimes
Quebec
Ontario
Manitoba
Saskatchewan
Alberta
British Columbia
12,567
171
368
339
1278
4681
5628
90
11,678
170
340
292
603
4969
5232
72
9573
113
302
291
1035
3397
4405
31
10,984
146
308
218
1195
3945
5114
58
11,781
123
258
192
1121
4594
5448
47
11,314
146
315
266
1122
4317
5165
60
100
1
3
2
10
38
46
<1
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Barley: Production, Improvement, and Uses
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fig. 8.2.1. Barley production regions in Canada: 1, Maritimes; 2, Quebec; 3, Ontario; 4, Southeastern Prairies; 5, South
Central Prairies; and 6, Northern Montane.
speciic characteristics and breeding priorities for
each region.
program in Ottawa has responsibility for this
area.
Québec region
Maritime region
This region is characterized by abundant moisture, high humidity, and moderate temperatures.
Acid soils are typical for this area. Foliar diseases
are common and lodging is often a problem.
There are no longer any barley breeding programs based in this area; thus, growers in the
region rely on germplasm developed elsewhere.
The Agriculture and Agri-Food Canada (AAFC)
F
The eastern portion of this region is similar to the
Maritimes, while the western portion has commonalities with Ontario. In both areas, resistance
to foliar diseases and strong straw are important
selection criteria. Breeding for this region is
carried out by the private sector since all of the
public breeding programs have been terminated.
The Ottawa program also has responsibility for
this area.
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Barley Breeding History, Progress, Objectives, and Technology
Ontario region
This region also is favored by adequate moisture;
however, temperatures are considerably higher
and midsummer drought is often encountered.
Thus, in addition to disease resistance and straw
strength, early maturity is desirable to avoid the
hot dry periods of July and August. A small area
of winter barley is produced in the southwestern
portion of this province. This region is serviced
by breeding programs at AAFC’s Ottawa station,
the private sector, and a small program at the
University of Guelph.
Western Canada
Western Canada is a vast area comparable in size
to continental Europe. Clearly, such a large area
has many ecozones contained within its borders.
It can, however, be divided into three broad areas:
the warm, moist Southeastern Prairies region; the
cool Northern Montane region; and the drier
South Central Prairies region.
Southeastern Prairies
The Southeastern Prairies region is characterized
by adequate moisture levels, in most years, high
temperatures and humidity in midsummer.
Diseases such as stem rust (incited by Puccinia
graminis f. sp. tritici [Eriks. and E. Henn.] D.M.
Henderson), fusarium head blight (FHB, incited
by Fusarium graminearum Schwabe), and spot
blotch (incited by Cochliobolus sativus [Ito and
Kuribayashi] Drechs. ex Dastur) are favored by
such conditions. The AAFC’s Brandon programs
play a key role in servicing this area. The
University of Saskatchewan’s Crop Development
Centre also provides cultivars for this area.
Cultivars developed for Minnesota and North
Dakota also do well here.
171
Drechslera teres [Sacc.] Shoemaker f. teres and D.
teres f. maculata Smedeg.) is the most frequently
found. This area is less suited to six-row cultivars
and short-statured cultivars for those producers
who windrow their crop prior to combining. This
is prime two-row malting barley production territory. This area is the prime focus of the Crop
Development Centre in Saskatoon but is also serviced by the Brandon and Lacombe programs.
Northern Montane
The Northern Montane region typically also has
adequate moisture but has cooler temperatures
and shorter growing seasons. Scald (incited by
Rhynchosporium secalis [Oudem.] J.J. Davis) is a
serious leaf disease in this region. Lower temperatures generally do not favor spot blotch and FHB.
The Alberta Field Crop Development Centre, in
Lacombe, is strategically located to service this
area. The area is also serviced by the programs in
Brandon and Saskatoon.
Of the 11–12 million tons of barley produced in
Canada each year, over 90% is grown in Western
Canada. Approximately 80% is used for feed,
most of that domestically. Canada’s exports of
barley are typically in the range of 2–3 million
tons. The largest portion of this is malting barley.
Major customers for malting barley are the
United States, China, Colombia, and South
Africa. United States and Japan are major importers of Canadian malt, amounting to several
hundred thousand tons annually. Markets for
feed barley are more variable with Saudi Arabia,
Japan, and several Middle Eastern countries as
the most consistent buyers. Food barley consumes a very small but growing amount of barley
and it constitutes an important niche for some
growers. Detailed statistics can be obtained from
the Canadian Grain Commission reports (http://
www.grainscanada.gc.ca/; veriied July 22, 2009). 23
BARLEY PRODUCTiOn in THE
UniTED sTATEs
South Central Prairies
The South Central Prairies region is characterized
as having reduced rainfall, high evapotranspiration rates, and frequent droughts. Foliar diseases
are less common and net blotch (incited by
Barley was introduced into the United States
along the Eastern Coast shortly after the irst
settlers arrived in the seventeenth century and
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Barley: Production, Improvement, and Uses
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its production moved west with the growth of
24 the population (Poehlman 1985). One factor
hastening the movement of barley to the west
was the disease FHB. This pathogen attacks not
only barley but also wheat (Triticum spp.) and
maize (Zea mays L.). As maize production would
move into an area, the level of FHB in the
barley would increase; thus, its production would
move west to areas where maize was not grown.
This movement of barley west to avoid FHB
continues today. The Red River Valley region
of North Dakota and Minnesota was the center
of six-row barley production in the United States
from the 1940s to the 1990s. In the 1990s, aboveaverage precipitation and an increase in maize
production in the Red River Valley provided
the conditions conducive to FHB in barley. Thus,
many growers in the region quit growing barley
because of the high susceptibility of the crop
to FHB and other market factors. The center
of six-row barley production in the United States
is now north central North Dakota, and new
cultivars are being developed that are adapted
to the more arid regions of western North Dakota
and eastern Montana.
The area of barley production in the United
States can be subdivided into four regions: the
East, Upper Midwest, West, and Southwest. The
Midwest and West regions account for over
80% of the area sown to barley (United States
Department of Agriculture National Agricultural
Statistics Service [USDA-NASS]; available at
http://www.nass.usda.gov/index.asp; veriied
May 23, 2009). Speciics mentioned below on
barley production and cultivars grown in the
United States and in many states can be obtained
from the Web site of the USDA-NASS (available
at http://www.nass.usda.gov/index.asp; veriied
May 23, 2009).
North Dakota is the largest barley-producing
state in the United States followed by Idaho and
Montana (USDA-NASS). Over 90% of the
barley sown in the Upper Midwest region of
Minnesota, North Dakota, and South Dakota are
six-row malting barley cultivars. Production in
this region is typically done under dryland conditions, and the reason for the preponderance of
six-row barley goes back to the origin of the barley
F
improvement efforts in the early 1900s. The
barleys found to be best adapted to the region and
to have acceptable malt quality at the time were
six-row barley introductions that could be traced
back to the Manchuria region of China (Rasmusson
1985).
In the West region, two-row barley cultivars
are typically grown, and barley for malting is
often grown under irrigation. Of the early barley
introduced into much of this region, two-row
barley accessions from central Europe were best
adapted and formed the germplasm pool from
which many malting barley cultivars were developed. The area sown to malting versus nonmalting types varies by state. Idaho and Montana, the
two largest barley-producing states in the region,
and Wyoming sow between 65% and 80% of
their barley area to malting cultivars. Other states
in the region may sow less than 10% of their
barley area to malting cultivars.
In the East region, nonmalting winter barley
cultivars are typically grown. Maryland,
Pennsylvania, and Virginia sow the most barley
in the region; yet, their combined production area
accounts on average for only 3.6% (5-year average,
2004–2008) of the U.S. total. In the Southwest
region, nonmalting spring barley is typically
grown and most of the production is in the irrigated desert regions of Arizona and Southern
California. The estimated total area sown to
barley in this region is around 1% of the U.S.
total.
Unlike Canada, most of the barley produced in
the United States is used domestically. From
2004 to 2007, less than 15% of the annual U.S.
production of 4.8 million tons is exported, and
most of the barley exported is feed barley (http://
usda.mannlib.cornell.edu/usda/fas/grainmarket//2000s/2008/grain-market-1211-2008.pdf and http://www.nass.usda.gov/
QuickStats/index2.jsp; veriied July 22, 2009).
BREEDinG PROGRAMs in CAnADA
Early barley improvement was carried out
through introductions, initially from Europe and
later from Manchuria. Selection carried out on
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Barley Breeding History, Progress, Objectives, and Technology
these Manchurian six-row landraces, irst at
Ontario Agricultural College in Guelph and later
at Macdonald College in Québec, gave rise to
important cultivars in Canadian barley history.
Most notable of these was OAC 21, which was
widely grown throughout both Eastern and
Western Canada. It was the standard for malting
quality for six-row barley for many decades.
Successful hybridization did not start until the
1920s; among the irst products of this program
from Macdonald College was Montcalm. This
cultivar was the irst Canadian malting barley
developed by hybridization. It was widely adapted
and high yielding and had excellent malting
quality for its time. Not surprisingly, it soon
dominated the malting barley acreage and held
that position until the mid-1960s.
In Western Canada, agricultural colleges were
established in all three Prairie Provinces by 1918.
As in the East, initial focus was on the introduction of cultivars from other sources. OAC
21, for example, was widely distributed by these
colleges. Hybridization programs were quickly
established, however, aimed at producing cultivars suitable for prairie conditions. The AAFC
research stations also became active at this time.
Notable among these was the station at Brandon,
Manitoba, a program that has remained extremely
important to this day.
Barley breeding in Canada is primarily carried
out by public sector agencies due to the lack
of proitability of private sector breeding in
self-pollinated crops like barley. Typically, less
than 10% of the barley acreage is sown with
pedigreed seed; thus, royalty collection is insuficient to generate proitability. These public
programs have a long history of collaboration
with each other and with programs in the United
States. A number of annual and periodic meetings within Canada provide networking opportunities for Canadian barley scientists. The
triennial North American Barley Researchers
Workshop is a venue in which barley scientists
from both countries exchange views and information, setting the stage for ongoing shared
targets and collaboration. Speciics on the public
breeding programs in Canada are presented
below.
173
AAfC, Ottawa
This program originally served eastern Ontario
and western Quebec but has expanded its original
mandate area to include the Maritime Provinces,
Quebec, and Ontario where programs have been
eliminated or reduced in scope. This is an enormous geographic area even though it only constitutes about 6% of the Canadian barley production.
Production here is used exclusively for feed
except for very small amounts of malting barley
in Quebec. Thus, the program has the daunting
task of servicing a diversity of production conditions. In this feed market, no premiums are paid
for quality, and thus the emphasis is on yield and
yield stability through disease resistance, straw
strength, and nutrient use eficiency. Important
diseases here include powdery mildew (incited by
Erysiphe graminis DC. f. sp. hordei Em. Marchal),
scald, FHB, stem rust, net blotch, Septoria speckled leaf blotch (incited by Septoria passerinii Sacc. 25
and Stagonospora avenae f. sp. triticea T. Johnson),
and barley yellow dwarf virus. This program and
its predecessors have provided excellent varieties,
primarily of six-row cultivars, to service the needs
of producers in this region.
University of Guelph
As indicated earlier, this program was one of the
original barley improvement programs in Canada
and provided cultivars like OAC 21, which gave
Canada wide importance for many years. This 26
program, in addition to cultivar improvement,
developed the Hordeum bulbosum method of
doubled-haploid barley production that had
widespread use in barley genetics research (Kasha
and Kao 1970). The program now is the responsibility of one breeder for a portion of his time.
The focus has narrowed to primarily six-row,
spring, feed types. Objectives are high yield,
disease resistance, and early maturity. Despite the
reduction in resources, this program continues to
play an important role for southwestern Ontario
producers.
AAfC Brandon
The AAFC program in Brandon, Manitoba, is
another program that has been historically
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Barley: Production, Improvement, and Uses
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important to barley production in Western
Canada. This program has an excellent record
of producing widely adapted six-row cultivars
for both feed and malting uses. Conquest and
Bonanza are the best known of these cultivars
and dominated the six-row malting area for
many years. This station also provided valuable
service by incorporating sources of resistance
for a number of diseases into adapted materials.
More recently, this role has expanded to tworow barley lines, and the broadly grown AC
Metcalfe is an example of the important cultivars
emanating from Brandon.
Alberta Agriculture field Crop
Development Centre, Lacombe, AB
This program is the result of the evolution of
AAFC’s programs at Lethbridge, Beaverlodge,
and Lacombe and at the University of Alberta, all
of which have been terminated (except Lacombe).
Soon after its inception, a federal provincial
agreement was signed and resources from both
governments were contributed. The vast majority
of its efforts have been devoted to developing
high yield potential cultivars for Alberta’s hog
and cattle industries. A number of cultivars have
been released with good disease resistance and
adaptation to high manure rates and adequate
rainfall.
University of saskatchewan
Since its inception during the early part of the
twentieth century, this has been the only
Saskatchewan-based breeding program. Its
primary focus has been the dryer portions of
the prairies. The program was given a signiicant boost with the establishment of the Crop
Development Centre in 1971. This recognized
the importance of barley in the province where
the largest acreage and the second largest production in Canada exists. The program has
been preeminent in the production of highquality two-row malting barley as typiied by
Harrington and a number of its successors.
Harrington set the standard for quality in high
enzyme activity, rapid modiication barley cultivars. It was also highly suited to the harsh
environment of the dry prairies. The feed and
food portion of the program pioneered a
number of barley utilization areas in Canada
including hulless feed barley, both two- and
six-row; hulless waxy food barley; two-row
forage barley; semidwarf six-row feed barley;
pure amylopectin hulless, waxy food barley;
hulless barley with resistance to both surfaceborne and loose smut (incited by Ustilago tritici
[Pers.] Rostr.) for organic production; hulless
malting barley; and low phytate barley. The
program is one of the few programs to date to
utilize molecular markers routinely for breeding, especially disease resistance, and for puriication of breeder seed.
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supporting facilities
In addition to the above breeding programs, there
are several units that are important contributors
to the breeding programs. Pathology support is
provided by AAFC’s scientists with the lead role
taken by the station in Winnipeg and support
from Lacombe, Charlottetown, and Ottawa.
These scientists have developed testing protocols
for all of the important barley diseases in Canada
and supervise and collaborate in screening for
resistance. This is of course a vital resource for
breeders. Plant Genetic Resources Canada, with
headquarters in Saskatoon, contains Canada’s
gene bank. Canada has global responsibility for
barley and oat, and thus has a large collection of
cultivated and wild species of barley. There is
excellent collaboration between breeders and
gene bank scientists in the collection characterization and utilization of barley germplasm.
Malting barley is an important cash crop for
Canada, and thus malting cultivars are sown on
far more area than end usage would justify. Given
Canada’s signiicant role in the export market for
malting barley and malt, the development of
high-quality potential cultivars is essential.
Malting barley breeders have their own screening
laboratories but rely on others for screening
advanced materials. This role is carried by several
organizations. The Grain Research Laboratory of
the Canadian Grain Commission develops testing
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Barley Breeding History, Progress, Objectives, and Technology
protocols, screens advanced lines from all programs, and collates the data for cultivar registration testing evaluation. Members of the industry
also contribute to this testing though their own
laboratories. The Brewing and Malting Barley
Research Institute (BMBRI) is an industry-based
organization that coordinates collaborative testing
of candidate malting lines for registration in
Canada. They also assist in arranging plantscale testing post registration. The Canadian
Malting Barley Technical Centre is an independent membership-based facility that provides
pilot-scale malting and brewing tests on a fee for
service basis. The Canadian Wheat Board is the
marketing agency for Canadian malting barley
and facilitates plant-scale testing of newly registered cultivars before inal acceptance in the
domestic and export markets.
All of these agencies located at Winnipeg collaborate to provide quality targets for breeders
that can be accessed at the BMBRI Web site
(http://www.bmbri.ca/; veriied May 23, 2009).
These are similar to those published by the
American Malting Barley Association (AMBA) in
the United States and are listed elsewhere in this
chapter.
There are unfortunately no similar networks
for feed and food barley; thus, breeders are forced
to set up any necessary testing on a case-by-case
basis.
BREEDinG PROGRAMs in
THE UniTED sTATEs
Barley breeding in the United States is largely
done by public institutions located in the northern tier of states. Institutions with breeders
working solely on barley include Montana State
University, North Dakota State University
(NDSU), Oregon State University, the University
of California, Davis, the University of Minnesota,
Washington State University, and the United
States Department of Agriculture Agricultural
Research Service (USDA-ARS) at Aberdeen,
Idaho. Public programs with breeders working
on barley as well as other small grains are located
at the University of Georgia, University of
175
Maryland, University of Nebraska, Utah State
University, and Virginia Polytechnic Institute
and State University (VPI). All of these programs
have development of wheat (Triticum aestivum
L.) cultivars as their top priority and have
much smaller efforts in barley. The programs in
California, Minnesota, Montana, North Dakota,
and Washington work primarily on developing
spring barley cultivars; the programs in Georgia,
Maryland, Nebraska, Oregon, and Virginia work
almost exclusively in developing winter barley
cultivars; and the USDA-ARS program in Idaho
works on developing both spring and winter
barley cultivars. In addition, the Minnesota,
North Dakota, and Idaho USDA-ARS programs
work primarily on developing malting barley cultivars. Unlike many areas of the world where only
malt made from two-row barley is used for
brewing, the brewing industry in the United
States uses substantial amounts of malt made
from six-row barley. The breeding program at the
University of Minnesota focuses on the development of six-row barley cultivars, whereas the
NDSU and USDA-ARS programs work on
the development of two- and six-row cultivars.
The public programs in California, Georgia,
Maryland, Nebraska, Utah, and Virginia develop
primarily nonmalting barley cultivars, and the
Montana, Oregon, and Washington programs
work on barley for malting and nonmalting
uses. The VPI, Idaho USDA-ARS, Oregon, and
Washington breeding programs also work on
hulless barley for food, feed, and industrial uses.
The program at VPI is particularly working on
hulless barley for ethanol production in collaboration with the USDA-ARS Eastern Regional
Research Center at Wyndmoor, Pennsylvania.
Breeding in the private sector is dominated by
three companies that develop cultivars for production across more than one barley-growing
region in the United States. The companies and
the headquarters of their breeding programs are
Busch Agricultural Resources, LLC (BAR, a
member of Anheuser-Busch InBev), located in
Fort Collins, Colorado; MillerCoors, located in
Burley, Idaho; and Westbred, LLC, headquartered in Bozeman, Montana. The BAR and
MillerCoors programs breed malting barley,
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Barley: Production, Improvement, and Uses
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whereas Westbred develops feed, hulless food,
and forage-type barley cultivars. The MillerCoors
program develops cultivars almost exclusively for
use within their company, whereas BAR develops
cultivars that are used by brewers in addition to
Anheuser-Busch InBev.
Breeding priorities in the United states
Malting
The choice of parents to use for crossing is critical
for developing acceptable barley cultivars because
barley cultivars often are adapted to speciic production regions and are designed for particular
uses, such as malting and brewing. Designation
of a cultivar as a “malting-type” is assigned by a
particular company that would use the cultivar or
by an organization such as AMBA, located in
Milwaukee, Wisconsin. AMBA is a nonproit
trade association composed of dues paying malting
and brewing companies. A primary goal of AMBA
is to ensure an adequate supply of high-quality
two- and six-row malting barley for the malting
and brewing industries in the United States
(http://www.ambainc.org/about/bh.htm; veriied May 23, 2009). This is accomplished, in part,
by providing grant funds to public breeding programs to support their cultivar development
efforts. AMBA also coordinates pilot-scale and
plant-scale evaluation programs of advanced
breeding lines or cultivars to determine their suitability for their member companies. These two
programs will be discussed in greater detail later
in the chapter. Evaluation of malt quality of
breeding lines prior to entering into AMBA’s
testing programs is done by the USDA-ARS
Cereal Crops Research Unit in Madison,
Wisconsin, and in limited cases in industry laboratories. Evaluation of breeding lines developed
by the public breeding programs at a single site
by the USDA-ARS allows AMBA’s members
and other stakeholders to make comparisons
between lines that are all malted and evaluated
under similar conditions.
Malting barley in the United States is unique
from most other crops in the country in that it is
marketed and stored on an identity preserved
basis. Malting barley cultivars are kept segregated
F
by cultivar, and often times a cultivar is additionally segregated based on production area or by a
speciic quality trait such as grain protein level.
Segregation is done because cultivars used for
malting and brewing must meet a long list of
speciic criteria before they are accepted. In addition, malting is done by cultivar and often by
cultivar grown at a speciic location, and many
beer brands have speciic compositions of different barley cultivars in their brewing blend. A
unique feature of malting barley in North America
is that the life of cultivars can easily be 10 years
or more. Examples of this are Larker, Morex,
and Robust six-row barleys, and Klages and
Harrington two-row barleys. Larker, Morex,
Robust, and Klages were developed in the United
States, and Harrington was developed at the
University of Saskatchewan in Saskatoon, Canada.
Guidelines for speciic quality traits, which
include measurements on barley and malt, are
provided to the public breeding programs by
AMBA (Table 8.2.2). Quality traits that receive
the most attention in malting barley breeding programs are barley grain protein and kernel plumpness, and malt extract, enzymatic activity
(α-amylase and diastatic power), wort protein,
measures of carbohydrate modiication (β-glucan
content and wort viscosity), and protein modiication (Kohlbach index) in the malt. A detailed
description of these malt parameters and others
can be found in Kunze (2004).
Because of the long list of parameters that must
be met, development of malting barley cultivars
is typically done using crosses between parents
that already have acceptable quality. Shortfalls in
malt quality of either parent usually appear in the
progeny. This need to make crosses between
parents with acceptable quality has led to very
narrow germplasm bases because variability
already has been severely restricted by adaptation
to speciic production regions. For example,
Horsley et al. (1995) stated that in the early 1990s,
all cultivars developed by six-row barley breeding
programs in the Upper Midwest United States
and the eastern Prairie Provinces of Canada could
be traced back to 15 accessions obtained in the
early 1890s. However, even within the narrow
germplasm bases of malt barley programs, gains
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Barley Breeding History, Progress, Objectives, and Technology
Table 8.2.2 Barley and malt quality speciications provided
to barley breeders in the United States by the American
Malting Barley Association, Inc., Milwaukee, Wisconsin,
in May 2008 (http://www.ambainc.org/ni/Guidelines.pdf;
veriied 22 July 2009)
135
Barley factors
Plump kernelsa
Thin kernelsb
Germinationc
Protein
Skinned and broken
kernels
Malt factors
Total protein
On 7/64” sieve
Two-Row Barley
Six-Row Barley
>90%
<3%
>98%
≤13%
<5%
>80%
<3%
>98%
≤13.5%
<5%
≤12.8%
>70%
≤13.3%
>60%
Measures of malt modiication
β-Glucan (ppm)
<100
Fine–coarse extract
<1.2
difference
Kohlbach index
40%–47%
Turbity (NTU)
<10
Viscosity (absolute
<1.50
cP)
136
Congress wort
Soluble protein
Extract (ine grind
db)
Color (°ASBC)
Free amino nitrogen
Malt enzymes
Diastatic power
(°ASBC)
α-amylase (20° DU)
<120
<1.2
42%–47%
<10
1.50
4.4%–5.6%
>81.0%
5.2%–5.7%
>79.0%
1.6–2.5
>190
1.8–2.5
>200
>120
>140
>50
>50
177
malting barley. However, feed quality attributes
are being developed for several classes of livestock. Using marker-assisted selection (MAS),
cv. Valier was developed to incorporate excellent
agronomic performance and improved feed
characters for beef cattle (Blake et al. 2002).
Incorporating disease resistance genes becomes
especially important because application of
expensive fungicides to feed barley can be cost
prohibitive.
Breeders developing hulless barley for human
consumption look to develop cultivars that are
high in soluble iber to reduce the risk of cardiovascular disease (Behall et al. 2004). The trait that
receives the most attention is breeding for
increased grain β-glucan content. Breeders developing hulless barley for the poultry feed market
work to develop cultivars with reduced grain βglucan content (Newman et al. 1987). Unlike the
central USDA-ARS-CCRU laboratory available 27
for testing malt quality, there is no central laboratory available for determining the feed or food
quality of breeding lines. Each of the breeding
programs is responsible for determining the
quality of their lines themselves. Likewise, there
is no single organization such as AMBA that is
providing desired quality characteristics to breeders for feed and food barley.
Resistance to diseases
still are being made. Rasmusson and Phillips
(1997) theorized that gains are made in the narrow
germplasm bases due to de novo variation and
elevated epistasis. In addition, for an excellent
review on gains and diversiication in barley
obtained through breeding, the reader is encouraged to read Fischbeck (2003).
Feed and food
In developing cultivars designed for livestock
feed, the choice of parents is of less importance
because growers are usually not paid premiums
for speciic quality parameters as is done for
A high priority in many breeding programs is the
development of cultivars with resistance to one or
more pathogens. Because barley is considered a
low-input crop, the easiest and most cost-effective
method of disease control is growing resistant cultivars. The list of barley diseases caused by bacteria, fungi, and viruses is plentiful (Mathre
1997). Good sources of genes for resistance to
diseases and other pests are often unadapted
accessions or the wild ancestors of barley, such as
Hordeum vulgare subsp. spontaneum. The USDAARS National Small Grains Collection (NSGC)
in Aberdeen, Idaho, is the part of the USDAARS’s National Plant Germplasm System that
has a large collection of cultivated and wild ancestors and relatives of barley and other small grain
cereals from around the world. In June 2009, the
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Barley: Production, Improvement, and Uses
178
USDA-ARS-NSGC had 27,925 accessions of H.
vulgare subsp. vulgare, 1507 accessions of H.
vulgare subsp. spontaneum, and over 440 additional Hordeum accessions (http://www.arsgrin.gov/cgi-bin/npgs/html/site_holding.pl?NSGC;
veriied July 22, 2009).
Another important utility that the USDAARS provides to barley breeders, pathologists,
and other stakeholders is tracking and identiication of different rust pathogens in the United
States, and screening and identiication of new
sources of resistance to new rust pathotypes. This
role is overseen by the USDA-ARS Cereal
Disease Laboratory (CDL) located in St. Paul,
Minnesota. Each year, the USDA-ARS-CDL
publishes a series of bulletins that report on the
current situation of several cereal rust diseases in
the United States (http://www.ars.usda.gov/
Main/docs.htm?docid=9757; veriied July 22,
2009).
Each barley-growing region of the United
States has different disease proiles and priorities
when breeding for disease resistance. In the
Upper Midwest United States, the high-priority
prevalent diseases are wheat stem rust, FHB,
spot blotch, and the net form of net blotch
(incited by Pyrenophora teres Drechs.). All cultivars developed for this region must have resistance to the prevalent pathotypes of wheat stem
rust due to the devastation this disease can cause.
Since 1993, major breeding efforts have been
ongoing in the region to develop cultivars with
improved resistance to FHB and accumulation of
the mycotoxin deoxynivalenol (DON) that is
produced by the causal organism. The best
sources of resistance to FHB and DON accumulation being used by Midwestern U.S. breeding
programs have originated from China, Japan,
and Switzerland (Prom et al. 1996; Urrea et al.
2005).
The list of foliar diseases causing signiicant
losses in the western and southwestern United
States is much smaller than that in the other
regions due to the much drier environment. The
disease receiving the most attention since the late
1980s is stripe rust, incited by Puccinia striiformis
f. sp. tritici. In 1975, a pathotype capable of attacking most cultivars in the United States spread
F
from Europe to Colombia, and the irst disease
caused by this pathotype was irst seen in the
United States in winter 1991 in Texas (Roelfs et
al. 1992). From Texas, the disease spread quickly
to California, Idaho, Montana, Oregon, and
Washington (Chen 2004). Because breeders and
pathologists were aware that this disease would
eventually reach the United States, they were
proactive and were able to identify and incorporate resistance into their germplasm (Roelfs and
Huerto-Espino 1994; Chen 2004). Complexes of 28
diseases that are more problematic in the western
United States are a series of different root and
crown rots (e.g., rhizoctonia, pythium, and fusarium). The effects of this group of diseases are
especially noticed in reduced tillage and direct
seed systems as well as during dry years or in dry
areas of the ield such as hilltops because roots are
unable to uptake suficient water (Cook and 29
Veseth 1991).
In the eastern United States, the predominant
diseases are barley leaf rust (incited by Puccinia
hordei Otth.), powdery mildew, and barley yellow
dwarf virus. A dificulty in breeding for barley
leaf rust is that multiple leaf rust resistance genes
have been overcome by changes in pathogen races
of P. hordei.
Resistance to insects and nematodes
In the United States, the aphids bird cherry-oat
aphid (Rhopalosiphum padi [L.]), green bug
(Schizaphis graminum [Rondani]), and Russian
wheat aphid (RWA) (Diuraphis noxia
[Kurdjumov]) are the major insect pests of barley
(Porter et al. 1999). Breeding for resistance to
RWA receives the most attention, and damage
from this pest occurs predominantly in the
intermountain west states of Colorado, Idaho,
Montana, and Wyoming. A large screening effort
for identiication of accessions in the USDAARS-NSGC with resistance to RWA was conducted in the glasshouse by USDA-ARS scientists
at Stillwater, Oklahoma (Webster et al. 1991).
Over 100 lines with RWA resistance were identiied and germplasm lines were rapidly developed
and made available to breeders (Mornhinweg
et al. 1995, 1999). In a collaborative effort
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Barley Breeding History, Progress, Objectives, and Technology
between USDA-ARS researchers at Stillwater,
Oklahoma, and Aberdeen, Idaho, additional
RWA germplasm lines and cultivars have been
developed. The cultivar Burton (Bregitzer et al.
2005) is an example of this successful collaboration of ARS scientists with university experiment
station researchers in the region.
Three
nematode
genera
(Heterodera,
Meloidogyne, and Pratylenchus) are known to
cause economic damage to barley (Mathre 1997).
However, breeding for resistance to nematodes in
the United States is not a priority.
Resistance to abiotic stresses
As stated earlier, barley is considered by growers
to be a low-input crop; thus, breeding for resistance to abiotic stresses is a high priority for many
barley breeders. A partial list of these stresses
includes drought and looding, high and low temperatures, mineral deiciencies and toxicities,
poor soil tilth (structure), and preharvest sprouting (PHS). A dificulty when breeding for abiotic
stresses is that plants often are exposed to multiple stresses at the same time, and it is not unusual
that common responses to the stresses are provoked in plants (Langridge et al. 2006). Thus,
breeding for abiotic stresses generally is considered more dificult than breeding for diseases or
other pests. Internationally, functional genomic
technologies are being used to gain a better understanding of how plants respond to abiotic stresses
(Langridge et al. 2006). However, establishment
of effective breeding strategies based on this
information will be challenging.
sELECTinG sOURCEs Of REsisTAnCE
TO ABiOTiC AnD BiOTiC sTREssEs in
CAnADA AnD THE UniTED sTATEs
After sometimes exhaustive searches, genotypes,
including cultivars, unadapted genotypes, landraces, and wild barley accessions that are resistant or tolerant to most economically important
biotic and abiotic stresses usually can be found
(Ullrich et al. 1995). However, sources of the
resistance genes are often poorly adapted to their
179
intended region, and desirable genes may show
linkage drag to undesirable traits. When a new
disease or pest arises or resistance breaks down
in currently grown cultivars, breeders can
develop improved cultivars in a short time if
they have been taking a proactive approach to
the problem. Examples of this approach include
work done by USDA-ARS researchers on RWA
and barley stripe rust beginning in the late 1980s.
The length of time needed to develop improved
cultivars also is reduced if the sources of resistance can be found in adapted germplasm. When
a loss of resistance is unanticipated, it can easily
take more than 10 years to incorporate new resistance into acceptable cultivars. In the case of
malting barley, this time frame is grossly underestimated because stringent quality parameters
must be maintained. This is the current case for
developing improved cultivars with resistance to
FHB and DON accumulation. Breeding for
these problems in the Midwest United States
began in 1993, and the irst breeding lines that
combined acceptable agronomic performance,
disease resistance, and malt quality did not enter
advanced testing by the malting and brewing
industries until 2008.
Not only is barley considered a low-input crop
by producers, but many barley breeders are
expected to develop improved cultivars using
limited inancial and physical resources. The tworow barley breeding program at NDSU in the
United States is an example where the existing
barley genetic resources were poorly adapted to
the production area when the program was started
in the early 1970s (Horsley et al. 2008). Improved
two-row lines were developed by making crosses
between unadapted two-row germplasm and
adapted Midwest U.S. six-row malting cultivars.
The most important selection criteria in early
generations were low grain protein (Foster et al.
1967) and high kernel plumpness. The irst cultivar released from this program was Bowman
(Franckowiak et al. 1985). Additional two-row
cultivars released by this program are Stark,
Logan, Conlon, Rawson, and Pinnacle. Conlon
was added to the list of cultivars recommended
for production as a malting barley by AMBA in
2000.
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Barley: Production, Improvement, and Uses
180
BREEDinG METHODs AnD
TECHniQUEs in CAnADA AnD THE
UniTED sTATEs
F
No two breeding programs use the same breeding
scheme. A detailed example of a breeding scheme
used to develop malting barley cultivars at NDSU
in the United States can be found in Horsley et
al. (2008). The scheme uses a modiied-pedigree
breeding method and off-season nurseries to
reduce the length of time needed to develop new
cultivars by up to 3 years. Variations to the scheme
above can be made at any stage to accommodate
speciic resources and breeding goals. For
example, doubled-haploids or MAS based on
DNA markers can be used to shorten the length
of time needed to develop new cultivars by several
years. These methods, as well as barley transformation, are described in greater detail in other
chapters. Off-season or glasshouse nurseries also
can reduce the length of time needed to develop
new cultivars by providing for two or more ield
generations per year. Locations used for offseason nurseries by breeders in Canada and the
United States include Arizona and California in
the southwestern United States, Argentina, and
New Zealand. Finally, nonmalting or feed barley
cultivars typically are developed as much as 3–4
years faster than malting barley cultivars because
30 extensive quality testing is not required.
Currently, MAS is gaining wider use and
research continues to identify opportunities to
incorporate this promising method. As stated
earlier, MAS is being used at the University of
Saskatchewan disease resistance breeding and
puriication of breeder seed. In the United States,
MAS is being used for identifying lines for resistance to such diseases as barley stripe rust, FHB,
and Septoria speckled leaf blotch. A 4-year (2006–
2010) multistate collaborative project in the
United States titled the Barley Cooperative
Agricultural Project (CAP), funded by the USDA
Cooperative State Research, Extension, and
Education Service (CSREES), has as its primary
goal to identify single nucleotide polymorphism
(SNP) markers associated with economically
important traits using association genetics
methods. Information from this project will be
used to generate PCR primers pairs for MAS at 31
the USDA-ARS high-throughput molecular
marker laboratories located in Fargo, North
Dakota, or Pullman, Washington. These laboratories are currently screening breeding lines using
molecular markers for traits such as FHB and
barley stripe rust resistance.
COLLABORATiOn BETWEEn BREEDinG
PROGRAMs in THE UniTED sTATEs
AnD AMBA fOR DEVELOPinG MALTinG
BARLEY CULTiVARs
A long history of collaboration has existed between
malting barley breeders in the Canadian and U.S.
breeding programs and members of the malting
and brewing industries. Development of improved
malting barley cultivars is dependent on barley
breeders knowing the desired levels of the numerous malt quality parameters required by maltsters
and brewers. Communication of the maltsters’
and brewers’ needs is conveyed to the breeders
through the Web and periodic meetings. As
described earlier, AMBA provides breeders in the
United States with baseline criteria that must be
met by new cultivars before they will be recommended for use by their members. This collaboration is accomplished through industry testing of
the malting and brewing qualities of advanced
breeding lines and new cultivars that is coordinated by AMBA. This evaluation, referred to as
pilot-scale and plant-scale evaluations, requires a
minimum of 4 years of testing. Before a cultivar
can be added to the “AMBA List of Recommended
Malting Barley Varieties,” it must be rated as
satisfactory in 2 years of pilot-scale evaluation and
in 2 years of plant-scale evaluation. Breeding programs participating in the AMBA evaluation programs include the public programs in Idaho,
Minnesota, Montana, North Dakota, Oregon,
and Washington, and BAR.
AMBA pilot-scale evaluation
Based on favorable agronomic and malt quality
characteristics, breeders are allowed to submit up
to eight lines (four six- and four two-row lines) to
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Barley Breeding History, Progress, Objectives, and Technology
AMBA for pilot-scale evaluation. Seed for this
evaluation is obtained from two of the best six
locations of large increase plots that are grown
speciically for AMBA’s tests by collaborating
breeders. For this evaluation, 17 kg of each entry
is divided among the malting and brewing
members of AMBA, and malt quality is compared
32 to the checked cultivars. No brewing evaluation
is done as part of the AMBA pilot-scale evaluation. Lines must be rated as satisfactory in 2 of 3
years of pilot-scale evaluation before they can be
considered for advancement to the AMBA Plant
Scale Evaluation Program.
181
the list in the 1980s (Azure, B1601, and Robust)
and 1990s (Excel, Foster, and Stander) for production in the Midwest United States. All six of
these cultivars are six-row. In the 2000s, seven
new cultivars (Conlon, Drummond, Lacey,
Legacy, Rasmusson, Stellar-ND, and Tradition)
have been added to the list for production in the
Midwest United States. Conlon is the only tworow cultivar. In addition, German cv. Scarlett was
added to the list in 2007 based on input from
AMBA’s malting members.
CULTiVAR RELEAsE in CAnADA
AMBA plant-scale evaluation
Typically, no more than one six-row and one tworow lines from a breeding program are allowed in
the plant-scale evaluation program at any one
time. The plant-scale evaluation program includes
evaluation of malting and brewing quality. Grain
of the candidate for plant-scale evaluation is
grown on approximately 250 ha in the region
where it is intended to be grown. The production
is done by local producers with the goal of producing a total 435–655 t of grain. Malt for plantscale brewing evaluation is typically produced by
one of AMBA’s malting members. In subsequent
years of the AMBA Plant Scale Evaluation
Program, a different maltster will be responsible
for producing the malt. This process ensures that
multiple maltsters have input during the AMBA
plant-scale evaluation process.
The methods of brewing evaluation of plantscale candidates are different for each brewing
company. Some companies may use small inclusions of the candidate line in their brewing blend,
while others may use inclusion rates of 50% or
more. Ultimately, each brewing member wants to
identify any deiciencies the candidate line may
have during the malting and brewing processes
and in the inished malt and beer. Even though
the addition of a cultivar to the AMBA List of
Recommended Malting Barley Varieties requires
support from only one of AMBA’s member companies, very few cultivars get added to the list
because of the stringent quality requirements.
For example, only three cultivars were added to
Cultivar registration is compulsory in Canada
for a number of ield crops including barley.
Registration is administered by the Variety
Registration Ofice of the Canadian Food
Inspection Agency (CFIA; http://www.
inspection.gc.ca/english/plaveg/variet/
vartoce.shtm; veriied July 22, 2009). This ofice
recognizes and relies on registration recommending committees to develop testing procedures, to
evaluate data, and to make recommendations on
the suitability of candidate cultivars for registration. In Western Canada, for example, the recognized committee is the Prairie Registration
Recommending Committee for Oat and Barley.
Details of testing protocols are posted on the Web
site http://www.pgdc.ca/ (veriied July 22,
2009). Membership on this committee includes
scientists from government, universities, and the
private sector; industry specialists; provincial
government specialists; and representatives of
farmer organizations. This committee sets the
standards for testing and tests are conducted
under its auspices. It also determines its operating
procedures subject to ratiication by the CFIA.
The committee is divided into three evaluation
teams for data evaluation purposes: the agronomy
team evaluates ield performance; the disease
team evaluates disease resistance; and the quality
team evaluates suitability for end-use processing
and manufacturing. The composition of these
yield trials, including checks, is approved by the
committee annually and administered by a test
coordinator. There are three yield trials: the 33
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Barley: Production, Improvement, and Uses
182
Western Six-Row Barley Cooperative, the
Western Two-Row Barley Cooperative, and the
Hulless Barley Cooperative. Entry into these tests
requires six station years of data from tests in
Canada with appropriate checks. Cooperative
preregistration trials are conducted by breeders
and other cooperators at a number of locations,
and candidates are evaluated over a 2-year period
before consideration for registration. Data are
collected on agronomic performance including
yield, lodging, maturity, seed weight, and so on.
In the case of malting barley, an additional year
of collaborative malting tests is required. Quality
evaluation is conducted on seeds from selected
sites of the yield trials. Micro malts are produced
and wet chemistry analyses are conducted by the
Grain Research Laboratory. In parallel to these
tests, a collaborative malting trial is coordinated
by the BMBRI. Promising entries from irst-year
cooperative testing plus successful entries from
previous collaborative testing are grown in separate plots, with commercial checks, at several sites
across Western Canada. Seed from selected sites
is malted and analyzed by the Grain Research
Laboratory and by industry laboratories. Disease
resistance data are collected on the yield trials
when infections justify and separate nurseries are
grown at various sites. For example, a stem rust
nursery is grown at the Winnipeg AAFC station;
a scald nursery is grown at Lacombe; and an FHB
nursery is grown at the Brandon AAFC station.
Coordinators are assigned for each disease and are
responsible for test coordination and data
collation.
When considering candidate cultivars, each
evaluation team passes expert judgment on their
area of expertise. A inal judgment is then made
by the committee in plenary on the basis of
all the data as a total package. Thus, these
experts now consider all data as opposed to
their respective discipline data so they are able
to balance weaknesses in one area with strengths
in another to reach a inal recommendation to
either support or not support the application
for registration. This information is transmitted
to the registration ofice and a inal decision is
made there, usually taking the advice of the
committee.
F
Postrelease adaptation and commercial
acceptance testing in Canada
Adaptation testing generates more information on
cultivar adaptation than is typically necessary for
a breeder to feel comfortable in releasing a new
cultivar and for cultivar registration purposes.
However, this information is found to be very
useful by many producers who use it to assist in
their decision to adopt new cultivars. The type of
testing in these yield trials is similar to advanced
generation yield trials but with a greater number
of stations involved and less data are recorded.
These trials are usually conducted by agencies
other than breeding organizations.
Commercial acceptance testing is necessary
for malting cultivars and for special use food
or feed cultivars. The malting and brewing
industry, for example, will not take on new
cultivars without such testing, in which they
support and actively participate. Typically, they
have seen several years of micromalting data on
new cultivars but no brewing information. Thus,
commercial scale ields are contracted and
selected barley is malted and brewed under commercial conditions. Most maltsters and brewers
want to see at least 2 years of successful malting
and brewing before they will begin to incorporate
a new cultivar into their blends. Again, these
tests, while critical to the success of a cultivar,
are beyond the purview of the breeder other
than to provide seed sources and to take a keen
interest in the process.
seed production in Canada
Pedigreed seed production in North America is
among the best in the world. The role of the
professional seed grower is vital to the success of
any new cultivar. They increase the seed that the
commercial farmer uses and maintain the genetic
integrity of the cultivar in doing so. The Canadian
Seed Growers Association (CSGA) and the CFIA
are coregulators of the pedigreed seed certiication system. The CSGA sets the standards and
the CFIA oversees production to ensure that
standards are met. For barley, there are the following classes of seed:
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Barley Breeding History, Progress, Objectives, and Technology
1.
2.
3.
4.
5.
Breeder seed, produced under the supervision
of a recognized breeder;
Select seed, produced from breeder seed by
qualiied select growers;
Foundation seed, produced by registered seed
growers;
Registered seed, and
Certiied seed.
The reader is referred to the CSGA Web site,
http://www.seedgrowers.ca (veriied July 22,
2009), for more detailed information.
CULTiVAR RELEAsE AnD inTELLECTUAL
PROPERTY issUEs in THE
UniTED sTATEs
Unlike many areas of the world, there is no formal
process in the United States administered by the
federal government for determining if a line can
be released as a named cultivar for commercial
production. Each university or private breeding
program has its own criteria and process for
releasing cultivars. While an experimental line is
usually evaluated in one or more Cooperative
Regional Yield Trials coordinated by the USDAARS, this is not required before it is released as a
named cultivar. The determination of whether a
cultivar will be added to the AMBA List of
Recommended Malting Barley Varieties is
described previously.
In general, most cultivars from the public programs are released to seed grower organizations
(e.g., Crop Improvement Associations) in their
states. The breeder and foundation seeds of these
releases are handled by the State Crop
Improvement Association or by the university or
state agricultural experiment station that released
the cultivar. Production of the registered and certiied seed classes is handled by growers often
contracted by seed companies, but ields and seed
are inspected and certiied by the State Crop
Improvement Association or State Department of
Agriculture. Production of the foundation seed
class for cultivars released by private companies
is not consistent. Some companies may contract
the production of the foundation class themselves
183
with local growers or they may contract a university to handle this production.
Protection of intellectual rights for public and
private breeders can be done by obtaining a plant
variety protection (PVP) certiicate administered
by the Plant Variety Protection Ofice of the
USDA Agricultural Marketing Service (http://
www.ams.usda.gov/AMSv1.0/; veriied July 28,
2009). Less commonly, utility patent protection
may be obtained through the United States Patent
and Trademark Ofice of the United States
Department of Commerce (http://www.uspto.
gov/main/patents.htm; veriied July 28, 2009).
The decision to obtain a PVP certiicate or a
utility patent for a new barley cultivar, especially
in the public sector, is not consistent across universities, agencies, programs, or even cultivars
nor is the institution of royalties for growing
protected cultivars.
REfEREnCEs
Behall, K.M., D.J. Scholield, and J.G. Hallfrisch. 2004.
Lipids signiicantly reduced by diet containing barley
compared to whole wheat and brown rice in moderately
hypercholesterolemic men. JAMA 23:55–62.
Blake, T., J.G.P. Bowman, P. Hensleigh, G. Kushna, G.
Carlson, L. Welty, J. Eckhoff, K. Kephart, D. Wichman,
and P.M. Hayes. 2002. Registration of ‘Valier’ barley.
Crop Sci. 42:1748–1749.
Bregitzer, P., D.W. Mornhinweg, R. Hammon, M. Stack,
D.D. Baltensperger, G.L. Hein, M.K. O’Neill, J.C.
Whitmore, and D.J. Fielder. 2005. Registration of Burton
barley. Crop Sci. 45:1166–1167.
Chen, X. 2004. Epidemiology of barley stripe rust and races
of Puccinia striiformis f. sp. hordei: the irst decade in the
United States. Cereal Rusts Powdery Mildews Bull. ••:••–
••. Available at http://www.crpmb.org/2004/1029chen/
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(accessed August 17, 2008).
Cook, R.J. and R.J. Veseth. 1991. Wheat Health Management.
APS Press, St. Paul, MN.
Fischbeck, G. 2003. Diversiication through breeding.
pp. 29–52. In R. von Bothmer, T. van Hintum, H. Knüpffer,
and K. Sato (eds.). Diversity in barley (Hordeum vulgare).
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Foster, A.E., G.A. Peterson, and U.J. Banasick. 1967.
Heritability of factors affecting malting quality of barley,
Hordeum vulgare L. Emend. Lam. Crop Sci. 7:611–613.
Franckowiak, J.D., A.E. Foster, V.D. Pederson, and R.E.
Pyler. 1985. Registration of Bowman barley. Crop Sci.
25:883.
Horsley, R.D., P.B. Schwarz, and J.J. Hammond. 1995.
Genetic diversity in malt quality of North American
six-row spring barley. Crop Sci. 35:113–118.
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Horsley, R.D., J.D. Franckowiak, and P.B. Schwarz. 2008.
Barley breeding. In M.J. Carena (ed.). Handbook of Plant
Breeding: Cereals, Vol. 3. Springer, Berlin (in press).
Jorgensen, J.H. 1992. Discovery, characterization and exploitation of Mlo powdery mildew resistance in barley.
Euphytica 63:141–152.
Kasha, K.J. and K.N. Kao. 1970. High frequency haploid
production in barley (Hordeum vulgare L.). Nature 225:
874–876.
Kunze, W. 2004. Technology Brewing and Malting, 3rd
international ed.—in English. VLB, Berlin.
Langridge, P., N. Paltridge, and G. Fincher. 2006. Functional
genomics of abiotic stress tolerance in cereals. Brief Funct.
Genom. Proteom. 4:343–354.
Mathre, D.E. (ed.). 1997. Compendium of Barley Diseases,
2nd ed. American Phytopathological Society, St. Paul,
MN.
Metcalfe, D.R. 1995. Barley. pp. 82–87. In A.E. Slinkard and
D.R. Knott (eds.). Harvest of Gold: the History of Field
Crop Breeding in Canada. University of Saskatchewan
Press, Saskatchewan, Canada.
Mornhinweg, D.W., D.R. Porter, and J.A. Webster. 1995.
Registration of STARS-9301B barley germplasm resistant
to Russian wheat aphid. Crop Sci. 35:603.
Mornhinweg, D.W., D.R. Porter, and J.A. Webster. 1999.
Registration of STARS-9577B Russian wheat aphid resistant barley germplasm. Crop Sci. 39:882–892.
Newman, R.K., C.W. Newman, and H. Graham. 1987.
Nutritional implication of β-glucans in barley. pp. 773–
780. In S. Yasuda and T. Konishi (eds.). Barley Genetics
V. Proc. 5th Int. Barley Genet. Symp., Okayama, Japan,
October 6–11, 1986.
Poehlman, J.M. 1985. Adaptation and distribution. pp. 1–17.
In D.C. Rasmusson (ed.). Barley. Agronomy Monograph
26. American Society of Agronomy, Madison, WI.
Porter, D.R., D.W. Mornhinweg, and J.A. Webster. 1999.
Insect resistance in barley germplasm. pp. 51–61. In S.L.
Clement and S.S. Quisenberry (eds.). Global Genetic
Resources for Insect-Resistant Crops. CRC Press, Boca
Raton, FL.
Prom, L.K., B.J. Steffenson, B. Salas, J. Moss, T.G. Fetch,
and H.H. Casper. 1996. Evaluation of selected barley
accessions for resistance to fusarium head blight and
deoxynivalenol concentration. pp. 764–766. In •• Scoles
and •• Rossnagel (eds.). Proc. of the V International Oat
Conference & VII International Barley Symposium,
University of Saskatchewan, Saskatoon, Canada. July 30–
August 6, 1996. University Extension Press, Saskatoon,
Canada.
Rasmusson, D.C. (ed.). 1985. Barley. Agronomy Monograph
26. ASA and CSSA, Madison, WI.
Rasmusson, D.C. and R.L. Phillips. 1997. Plant breeding
progress and genetic diversity from de novo variation and
elevated epistasis. Crop Sci. 37:303–310.
Roelfs, A.P. and J. Huerto-Espino. 1994. Seedling resistance
in Hordeum t barley stripe rust from Texas. Plant Dis.
78:1046–1049.
Roelfs, A.P., J. Huerta-Espino, and D. Marshall. 1992.
Barley stripe rust in Texas. Plant Dis. 76:538.
Ullrich, S.E., D.M. Wesenberg, H.E. Blockelman, and J.D.
Franckowiak. 1995. International cooperation in barley
germplasm activities. pp. 157–170. In R.R. Duncan (ed.).
International Germplasm Transfer: Past and Present.
CSSA and ASA, Madison, WI.
Urrea, C.A., R.D. Horsley, B.J. Steffenson, and P.B.
Schwarz. 2005. Agronomic characteristics, malt quality,
and disease resistance of barley germplasm lines with
partial fusarium head blight resistance. Crop Sci.
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Publications, Minneapolis, MN.
Webster, J.A., C.A. Baker, and D.R. Porter. 1991. Detection
and mechanisms of Russian wheat aphid (Homoptera:
Aphidadae) resistance in barley. J. Econ. Entomol.
842(2):669–673.
AUSTRALIA
David M.E. Poulsen and Reg C.M. Lance
HisTORY—BARLEY PRODUCTiOn
in AUsTRALiA
Barley is the second most important cereal crop,
after wheat, in Australia. It has expanded from
the irst crop of 3.24 ha, sown at Farm Cove after
the arrival of the First Fleet in 1788 (Sparrow and
Doolette 1975, 1987 loc. cit.), to an industry that 40
annually produces 4.5–5.5 million tons of grain
(Lovett 1997; Powell 1997; Stuart 1997; Healy
2001). Lovett (1997) stated that the net value of
Australian barley production was greater than
A$1 billion per year.
The barley-growing regions are conined to the
southern temperate cereal-growing regions with
rainfall varying from 250 to 600 mm (see Fig.
9.5.1, Chapter 9 Australia; Paynter and Fettell
2008) The igure indicates regional production as
tons per square kilometer based on the Australian
Bureau of Statistics 2006 Agricultural Census
data and is for the 2005/2006 harvest. The areas
of greatest production are southern Western
Australia; the southern Eyre Peninsula, Yorke
Peninsula, midnorth and upper southeast of
South Australia; the “Mallee” and Wimmera
regions of Victoria; southern New South Wales;
and the Darling Downs of Queensland. Other 41
production regions include northern New South
Wales and Tasmania.
Most Australian barley cultivars are two-row
spring types with white aleurone. In most areas,
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Barley Breeding History, Progress, Objectives, and Technology
the crops are sown in late autumn or early midwinter, lower in spring, and are harvested in
early summer.
Australian barley production over the 5-year
period 2002–2006 inclusive averaged 7 million
42 tons of which 2.619 MT was used domestically
and 4.590 MT was exported (Table 8.3.1). Only
0.169 MT was used for malt and other human
uses, and most (2.257 MT) are used for feed. Of
the exported grain, 2.461 MT was feed, 1.485 MT
was malting barley, and 0.627 MT was malt (in
grain equivalents). This malt is mostly exported
to Asian countries, vis-à-vis, Vietnam, Japan,
South Korea, the Philippines, and Thailand (Emalt.com 2008). So, in total, malt production was
0.796 MT. Although Australia is not a major
malting barley producer, its exports represent
approximately 25% of the world trade.
The largest barley production is in South
Australia (SA), closely followed by Western
Australia (WA), then Victoria (Vic), New South
Table 8.3.1 Australian barley production and disposal (5year average, 2002–2006)
Australian Barley Production
137 and Use
5-Year Average (2002–2006)
000 t
Production
7115
Domestic use
As malt and other human
use
Feed
Seed
2619
169
2257
193
Export
Feed barley
Malting barley
Malt (grain equivalent)
4590
2461
1485
627
Source: Australian Bureau of Agriculture and Resource
Economics (ABARE) and Australian Bureau of Statistics
(ABS).
Table 8.3.2
185
Wales (NSW), Queensland (Qld), and Tasmania
(Tas). Yields are highest in the southern regions 43
and lowest in New South Wales and Queensland
(Table 8.3.2).
For practicality of research management, the
Grains Research and Development Corporation
(GRDC) has divided the Australian cropping
environment into three regions, namely, the
northern region, the southern region, and the
western region (Davies 1993). The northern 44
region includes Queensland and northern New
South Wales. Southern New South Wales,
Victoria, South Australia, and Tasmania make up
the southern region. The dividing line between
the northern and southern regions is drawn at
Dubbo (latitude 33° S). Only Western Australian
cropping areas are included in the western region.
During the 1990s, approximately 63% of the
Australian barley crop was produced in the
southern region (Healy 2001). The western region
contributed 23% of the crop, while the northern
region produced approximately 14%. These
igures often vary, as seasonal conditions in
Australian barley-growing areas can luctuate
widely (Powell 1997). In most seasons, 40%–50%
of the Australian barley crop is accepted as malting
barley (Lovett 1997; Stuart 1997; MacLeod
2001). However, only 700,000 t or 35% of malt
barley production is malted within Australia
(MacLeod 2001). Approximately two-thirds of
the Australian malt production is exported annually (Powell 1997). Thus, approximately 88%
of the total Australian malting barley crop is
exported, making the country the third largest
exporter of malt, after Europe and North America
(MacLeod 2001). Consequently, the viability of
the Australian barley industry is heavily reliant on
maintaining its share of the international market
(Stuart 1997).
Australian barley production by state (5-year average, 2001–2005)
State
Units
Qld
NSW
Vic
Tas
SA
WA
Australia
Area
Production
Yield
Hectares (’000)
Tons (’000)
Tons per hectare
113
175
1.55
778
1356
1.74
793
1477
1.86
8
26
3.06
1176
2242
1.91
1160
2126
1.83
4030
7402
1.84
138
F
Source: Australian Bureau of Agriculture and Resource Economics (ABARE) and Australian Bureau of Statistics (ABS).
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Barley: Production, Improvement, and Uses
186
The main export market for Australian malting
barley is the Asia Paciic region. China buys
approximately 50% of the Australian malting
barley crop each year (Stuart 1997). Furthermore,
Sewell (1997) predicted that Chinese import
requirements of malting barley from Australia are
likely to reach 1.36 million tons by the year 2010.
This has arisen through increased demand for
beer within China and increased foreign investment in Chinese breweries (MacLeod 2001).
Because brewhouse technology has been reined,
quality speciications for the Chinese market have
become more stringent (Spiel 1999).
Japan is also a key market for Australian malting
barley and commands premium prices. In the
early 1980s, Australia’s share of the Japanese
malting barley market was approximately 33%
(Powell 1997). However, by 1997, signiicant
improvements in the quality of Canadian and
European cultivars led to Australian malting
barley exports to Japan being reduced to 124,226 t,
or 15.5% of Japan’s malt barley intake (Fukuda
et al. 1999). Other important markets for
Australian malting barley include Taiwan, the
Philippines, South Korea, and several other Asian
and South American countries (Stuart 1997).
Australian malt is principally exported to
Japan, Thailand, the Philippines, and South
Korea (MacLeod 2001). While the Japanese
market for Australian malt has declined in a
similar pattern to that seen for malting barley,
there have been signiicant increases in sales to
Thailand, Vietnam, and South Korea (E-malt.
com 2008). These three countries account for
47% of Australian malt exports. Increases in
brewing capacity in Thailand, Vietnam, and
South Korea should increase demand for
Australian malt and malting barley but will also
necessitate stable grain supplies and improved
variety performance.
In response to the above market signals, the
Australian industry has taken several steps to
improve the quality and reliability of its export
barley and malt supplies. The accreditation of
Australian malting cultivars is coordinated by the
Malting and Brewing Industry Barley Technical
Committee (MBIBTC) (Hawthorne 1999). This
committee has established technical guidelines,
F
which are available to all breeding programs.
Those guidelines include detailed experimental
schedules and speciications to be met by potential cultivars before domestic and/or export
malting accreditation is awarded (MBIBTC
1998).
Malting barley usually attracts a price premium
of A$30–A$50 per ton over feed barley in
Australia. To qualify for malting grade, the grain
must irstly be obtained from an accepted malting
cultivar. Point-of-delivery assessment must also
prove that the grain meets appropriate speciications, typically, a low proportion of small grain or
“screenings,” protein between 8.5% and 12.0%,
bright grain color, and absence of contaminants
such as smut, sclerotes, weed seeds, and soil (ABB
Grain 2008; CBH Group 2008; Grainco 2008). 45
HisTORY—BARLEY iMPROVEMEnT
in AUsTRALiA
The early history of Australian barley improvement was comprehensively described in review
papers written by Sparrow and Doolette (1975,
1987 loc. cit.). Most early Australian cultivars
were obtained from Europe. These tended to be
late maturing and befell the consequences of terminal droughts or the hot dry inishes to the
seasons so typical in southern Australia. A South
Australian farmer, Samuel Prior, developed the
irst Australian barley cultivar in 1903 from a
selection of imported seed line of Chevalier (or
alternatively, Spratt Archer). Prior’s Chevalier
selection was recognized for its superior malting
quality and subsequently proved to be well
adapted to the dry South Australian environment.
It became known as Prior and led to a massive
expansion in barley production. Prior became the
backbone of the Australian barley industry until
the end of the 1960s.
From the 1920s to the 1950s, barley breeding
was conducted on a part-time basis in Victoria,
New South Wales, and South Australia. Early
breeding programs led to Prior A bred by Albert
Pugsley and Research from the program at
Werribee in Victoria, bred by A.R. Raw in 1942.
However, in 1956, a barley improvement scheme
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Barley Breeding History, Progress, Objectives, and Technology
was established with funds provided by maltsters,
brewers, and growers and with matching funds
from the commonwealth government. This
scheme led to the development of full-time breeding programs in South Australia and Victoria, and
consequently, the release of several barley cultivars, which inally replaced Prior. The most signiicant of these was the malting barley Clipper,
released from the Waite Agricultural Research
Institute in 1968. The genotype × environment
interaction studies of Finlay and Wilkinson (1963)
had a signiicant inluence on the South Australian
program. Improved yield over a range of environments was found in introductions from North
Africa. This material was early maturing, tolerant
of alkaline and sodic soils, disease resistant (especially cereal cyst nematode [CCN], mildew, and
other leaf diseases), and tolerant of hot dry inishes. A similar well-adapted material was sourced
from the United States, but this originally came
from North Africa.
New breeding programs were established in
Western Australia in 1962 and in Queensland,
New South Wales, and Tasmania in the late 1960s
and early 1970s. Barley improvement in
Queensland effectively began in 1967 when Dr.
R.P. Johnston of the Department of Primary
Industries initiated a series of trials to evaluate
breeding lines from the Waite Institute (Poulsen
2001). A breeding program was designed to breed
high-yielding malting and feed barley cultivars,
targeting Queensland barley-producing regions.
Funding management for barley research and
development went through a series of iterations
from the late 1970s through to the present day
(Joppich 1985; Purse and McNee 1985; Smith
1987; Davies 1993). Originally, funding for different crops was governed through a series of
research councils. However, in 1990, 4 federal
research councils and 10 state wheat and barley
committees were amalgamated to form the
GRDC, with the overarching responsibility to
drive research and development across the entire
Australian grains industry (Davies 1993). This
has led to the development of a coordinated barley
research and development program for Australia,
including strong barley improvement programs
in each of the southern, western, and northern
187
regions (Lovett 2001). Further agronomic
research of French and Schultz (1984a,b) has
drawn attention to the importance of water use
eficiency in water-limiting environments and the
importance of understanding abiotic and biotic
constraints, which may limit productivity and
quality improvements.
Nationally focused projects have been established. For example, the National Barley
Molecular Marker Program (NBMMP), later to
be amalgamated with the equivalent wheat
program to become the Australian Winter Cereal
Molecular Marker Program (AWCMMP), was
set up to develop and implement molecular technologies on a national scale (Langridge 1997; Barr
et al. 2001).
The effects of developing focused barley breeding programs became clear during the 1980s, as
new malting cultivars with speciic regional adaptation were released, replacing Clipper (Sparrow
1984). Schooner was released from the Waite
Institute in 1983 and quickly dominated the
southern States. In Western Australia, the locally
bred cv. Stirling similarly took over from Clipper.
A third cultivar, Grimmett, replaced Clipper in
Queensland and in northern New South Wales.
The result of the signiicant investment from
the GRDC together with inputs from the malting
and brewing industries and from marketing and
grain handling organizations and authorities have 46
led to a signiicant improvement in the number
and quality of malting and feed varieties released
in the 1990s and early 2000s. Other cultivars have
since been released in each of the GRDC regions,
including high-yielding feed types.
A representative malting barley pedigree chart
is shown in Fig. 8.3.1, where possible varieties are
listed in chronological order. Caution should be
taken in attempting to determine the order of
each cross; special expediency sometimes prevented the convention of putting the female
parent irst. Some observations can be made.
Early programs were sparse and based on a limited
number of European parents. There was a limited
use of backcrossing to incorporate disease resistance traits. Cycle times were often 12 or more
years. The 1970s and 1980s saw the use of North
African landraces providing both an appropriate
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Barley: Production, Improvement, and Uses
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fig. 8.3.1. ••.
phenological adaptation and disease resistances
(such as CCN, mildew, and alkaline soil). New
introgressions came from Europe, North America,
and Japan. More varieties have been released with
shorter cycle times. The use of molecular genetic
markers has enabled more complex crosses to be
managed more easily and with greater eficiency
(e.g., Flagship, Sloop-SA, and Sloop-Vic). Barley
Australia accredited varieties are shown as shaded
rounded rectangles.
The newer malting varieties include Western
Australia: Gairdner, Baudin, Hamelin, and
Vlamingh; South Australia: Sloop and the backcross derivative Sloop-SA; Tasmania: Shannon,
Franklin, and Vertess; Queensland: Tallon and
Lindwall; Victoria: Arapilies, Sloop-Vic, Buloke,
F
and Hindmarsh; and New South Wales:
Cowabbie.
Feed varieties and some malting lines are
shown in Table 8.3.3. The table has been sorted
by date of release (or introduction) and by name.
RECEnT PROGREss in AUsTRALiAn
BARLEY iMPROVEMEnT PROGRAMs
Barley Breeding Australia (BBA)—
a new paradigm
BBA began operating on July 1, 2006 as the
Australian national barley breeding program
implementing a national plan for breeding
improved varieties to beneit the barley industry.
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Table 8.3.3
Australian feed barleys and other signiicant varieties
Name
Year Released
Pedigree
Breeder
Trabut
Abyssinian
Commander
Maltworthy
Deputy
Ablyn
Noyep
Anabee
Resibee
A14
A16
Bussell
Ketch
Lara
Cutter
Corvette
Parwan
Bandulla
Beecher
Cantala
Malebo
Waranga
O’Connor
Moondyne
AB6
Windich
Yerong
Gilbert
Brindabella
Kaputar
Namoi
Dash
Mundah
Molloy
Picola
Fitzgerald
Tilga
Unicorn
Doolup
Lindwall
Tantangara
Wyalong
Yambla
Keel
Binalong
MacKay
Cowabbie
Milby
Tulla
Capstan
Maritime
Grout
Urambie
Vertess
Yarra
Fleet
Hannan
Lockyer
Roe
1916
1920
1930
1938
1939
1952
1959
1962
1962
1968
1968
1968
1969
1971
1975
1976
1979
1981
1981
1981
1981
1981
1984
1987
1988
1989
1990
1992
1993
1993
1993
1995
1995
1996
1996
1997
1997
1997
1998
1998
1998
1998
1998
1999
2001
2001
2002
2002
2002
2004
2004
2005
2005
2005
2005
2006
2007
2007
2007
Algerian introduction selection
Abyssinian introduction
Coast selection
Prior × Beaven’s Special
Hero selection
Abyssinian × Flyn
Prior’s Chevalier selection
Research selection
Research selection
Prior × Ymer
Prior × Ymer
Prior × Ymer
Lenta × Noyep
Research × Lenta
Prior A × Proctor
Bonus × CI-3576
((((Plumage Archer × Prior) × Lenta)) × (Research × Lenta))
((Prior × Lenta) × (Noyep × Lenta))
Atlas × Vaughn
Kenia × Erectoides 16
Pallidum selection (CPI-11083)
((Plumage Archer × (Prior × Lenta)) × (Research × Lenta)) × Clipper
WI-2231 × (Atlas 57 × A14)
((Dampier × A14) × Kristina)) × (Clipper × Tenn 65-117)
CPI-71283 × 4* Clipper
(Atlas 57 × A16) × Parwan
M22 × Malebo
(Armelle × Lud) × Luke
((((Weeah × CI-17115) × HCB27) × Jadar II) × Cantalla)
((5604 × 1025)) × ((Emir × Shabet) × CM67)
(Sultan × Nackta) × (RM1508 × Godiva)
(Chad × Joline) × Cask
O’Connor × Yagan
(Golden Promise × WI-2395) × 72S:267
VB75031 × Elgina
Onslow × Tas 85-466
Forrest × Cantala
54C25 × 51C38
75S:323 × 74S:314
Triumph × Grimmett
AB6 × Skiff
Schooner × Stirling
Skiff × FM 437
(CPI-18197 × Clipper) × WI-2645
Blenheim × (Skiff × O’Connor)
Cameo × Koru
(((AB6 × Franklin) × Franklin)) × (Rubin × Skiff)
(((AB6 × Franklin) × Franklin)) × (Rubin × Skiff)
Skiff × FM 437
(Waveney × WI-2875) × (Chariot × Chebec)
74S:314 × 74S:309
Cameo × Arupo “S”
(Yagan × Ulandra) × Ulandra
Franklin × Cooper
(VB9108 × Alexis) × VB9104
Mundah × Keel × Barque
B28719 × (Windich × Morex)
Tantangara × VB9104
Doolup × (Windich × Morex)
DANSW
NSWDA
DANSW
RAC
DANSW
NSWDA
RAC
VicDA
VicDA
DAWA
DAWA
DAWA
WARI
VicDA
WARI
WARI
VicDA
VicDA
VicDA
NSWDA
Vic-DPI
DAWA
DAWA
CSIRO-PI
DAWA
NSWDPI
Qld-DPI
NSWDPI
CIMMYT
NSWDPI
NFC
DAWA
DAWA
Vic-DPI
DAWA
NSWDPI
Kirin Australia
DAWA
Qld-DPI
NSWDPI
NSWDPI
NSWDPI
WARI
NSWDPI
Qld-DPI
NSWDA
NSWDA
NSWDPI
Uni Adelaide
Uni Adelaide
Qld-DPI
NSWDPI
UTas and TASDPIWE
MBQIP
Uni Adelaide
DAFWA
DAFWA
DAFWA
F
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BBA is an unincorporated joint venture between
the GRDC, the Queensland Department of
Primary Industries and Fisheries, the New South
Wales Department of Primary Industries, the
Department of Primary Industries, Victoria, the
University of Adelaide, the Department of
Agriculture and Food, Western Australia, and the
Tasmanian Institute of Agricultural Research.
BBA is governed through an advisory board
formed by the parties and coordinated through a
management committee. The formation of BBA
rationalized the previous six state-based barley
breeding programs into one national program
with three regional nodes: BBA-North (QDPIWarwick, Queensland), BBA-South (University
of Adelaide, Urrbrae, South Australia), and BBAWest (DAFWA, Kensington, Western Australia).
The restructure aims to use resources clearly and
eficiently in barley breeding so as to provide the
foundation of a future vibrant barley industry
(Lance et al. 2007; Reading 2007). The previous
programs in Victoria, New South Wales, and
Tasmania will be focusing their efforts on germplasm development and barley prebreeding. The
breeding nodes are to focus on the development,
release, and adoption of varieties meeting speciic
market and regional needs. As such, they must
achieve operational eficiencies through the adoption of a “best practice in breeding” framework.
BBA-North is actively changing its primary
focus from breeding malt varieties to breeding
feed varieties, thereby meeting the future demand
for feed grain in northeastern Australia.
BBA-South has extended its trial network to
cover the neutral to alkaline soils in New South
Wales and Victoria and the alkaline soil regions of
southern Western Australia. The most adapted
barleys are mid season to earlier maturing.
BBA-West, in addition to conducting trials in
Western Australia, conducts yield trials and nurseries in New South Wales, Victoria, and Tasmania,
on neutral to acid soils. As such, the focus is on
medium- to later-maturing varieties suited to the
medium- to high-rainfall zones.
Germplasm from the NSWDPI, Victorian and
Tasmanian breeding programs is being pro47 gressed and integrated into the operations of
BBA-West and BBA-South.
Barley Australia—malting
barley accreditation
Barley Australia was established in 2005 as the
peak body for the Australian barley industry. It
was the recognition of a future deregulated marketing environment for Australian barley that was
a catalyst for its creation as it was recognized by
the maltsters and marketers (purchasing the
barley crop from the Australian farmers) that
there was a need for a body to deal with “industry
good” functions that had previously been undertaken by the statutory marketing companies that
preexisted the new market environment, such as
cooperative industry decisions on quality standards, variety suitability to malting, and a coordinated approach to industry-wide research.
Barley Australia is entirely self-funded by the
maltsters and marketers of Australian barley and
is a testament to the members in their ability to
cooperate for industry beneit even though in the
Australian marketplace they are staunch competitors. Now in its fourth year, Barley Australia
serves a signiicant purpose in the Australian
industry as a “shop window” for the industry and
disseminates information to the Australian barley
industry stakeholders as well as to the international marketplace. Barley Australia is responsible
for creating and managing the national varietal
accreditation system as well as the Barley Australia
Assured Quality trademark for HACCP-certiied 48
barley and malt out of Australia. More information on Barley Australia can be sought from the
Web site http://www.barleyaustralia.com.au/. 49
The MBiBTC
The MBIBTC has technical malting and brewing
experts representative of Australia’s major malting
and brewing companies. Varieties submitted to
Barley Australia are evaluated by an MBIBTC
member for the irst year of commercial evaluation. If deemed suitable, then the malt from the
evaluation passes to Pilot Brewing Australia
(PBA).
PBA
PBA conducts the last step in the process of evaluation of malting barley for either domestic or
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Barley Breeding History, Progress, Objectives, and Technology
export market suitability. PBA is malting and
brewing industry funded and also cofunded by
the GRDC. Samples for pilot brewing are taken
from commercial-scale malting evaluation trials.
The facility is able to replicate the requirements
of both domestic and export brewers utilizing
sugar or starch adjuncts.
For a new malting variety to be accredited, it
must pass two successful years of commercial
malting evaluation and pilot brewing or
commercial-scale brewing with all stages being
reviewed by the MBIBTC.
The major proportion of the northern region
barley crop is purchased for domestic stock feed.
Most northern feed barley is used for cattle since
the region contains the highest concentration of
50 feedlots in Australia (Barr and Kneipp 1995).
Feed for the monogastric industries is also a signiicant market, with approximately 30% of the
grain purchased by stock feed manufacturers for
pig and poultry rations (M. Covaceveitch, pers.
comm.).
AUsTRALiAn BARLEY BREEDinG
OBJECTiVEs
Barley breeding objectives have changed substantially in Australia over the past century. While the
basic agronomic objectives of grain yield, lodging
resistance, and appropriate maturity have changed
relatively little, the evolution of on-farm cultural
practices has resulted in a signiicant increase in
the number of diseases affecting the crop.
Objectives relating to grain quality have also
increased in number and detail, as knowledge of
grain quality for both malting and feed use has
become more reined. See Table 8.3.4 for a comprehensive summary of breeding objectives/
traits.
Agronomic traits
Yield, lodging resistance, straw strength, height
(short straw, medium height straw); phenology
(late maturity, short straw for longer seasons;
early maturity, taller straw for drier shorter
51 seasons); head loss, stem breakage.
191
Improved grain yield has been the primary
objective of Australian barley breeding activities
since the nineteenth century (Sparrow and
Doolette 1975). From the outset, modern
Australian programs were clearly designed with
yield as the primary selection criterion (Ellis
1983; Johnston 1983; Portman 1983; Read 1983;
Sparrow 1983; Vertigan 1983). However, it is
considered extremely important that yield
advances are made together with improvements
in grain quality to protect the Australian share of
the world barley market (GRDC 2001). Johnston
(1974) had concluded that a combined malting
and feed breeding program could service both
markets, improving both yield and quality.
As well as grain yield, Australian barley breeders have selected material for a range of agronomic traits. All of the programs made resistance
to lodging or a high level of straw strength a key
agronomic breeding objective (Portman 1983;
Read 1983; Vertigan 1983). Johnston (1983)
placed a very high priority on lodging resistance
as the inability of barley cultivars to stand up in
the ield was seen as a major limitation to the
further expansion of the crop in Queensland.
Plant height was established as another objective
of the Australian programs. For example, short
cultivars were desirable in the high- and mediumrainfall zones of Western Australia, while taller
cultivars suited low-rainfall areas (Portman 1983).
Controlling head loss caused by stem breakage
was another signiicant agronomic trait targeted,
in particular, for Western Australia, New South
Wales, and Tasmania.
Crop phenology targets of the different
Australian programs were diverse and were driven
by the amount of available moisture and the duration of optimal growing conditions. Read (1983)
explained how longer-season, short-straw material was being developed for parts of southern
New South Wales where early-sown cultivars
were suitable. These types were also suitable for
Tasmania and eastern Victoria. In Western
Australia, longer-season cultivars were more suitable for the high- and medium-rainfall zones,
while quick maturing barleys were required for
the parts of the cropping area with low rainfall
(Portman 1983). Most areas of South Australia
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Table 8.3.4 Differences in the grain production environments and key selection targets and traits for GRDC breeding
programs in northern, southern, and western regions
Southern Region
Western Region
Northern Region
Latitude 33° S–43° S
Predominantly winter rainfall
Lighter, shallower soils, sands in WA
Winter cropping only
Latitude 28° S–35° S
Predominantly winter rainfall
Lighter, shallower soils, sands in WA
Winter cropping only
Latitude 22° S–33° S
Predominantly summer rainfall
High proportion of heavy clay soil types
Winter/summer crop rotations
Double cropping
Agronomic traits
Yield, grain weight, plumpness, hectoliter weight, seedling vigor, phenological adaptation (bvp, ppd, vrn)
Malting quality
Malt extract, diastatic power (DP), apparent attenuation limit (AAL), low viscosity, increased free alpha amino nitrogen
(FAAN)
Feed quality
Digestible energy (DE), metabolizable energy (ME), animal intake, low bloat, low phytate
Resistance/tolerance to biotic stress
Net form net blotch
Spot form net blotch
Scald
Powdery mildew
Leaf rust (part of SA, S-NSW, NE,
and SW-Vic, Tas)
BYDV
Barley grass stripe rust
Rhizoctonia
139
Major pests
Cereal cyst nematode (SA, Vic)
Resistance/tolerance to abiotic stress
Drought/terminal drought
Alkaline soil/boron toxicity tolerance
(SA, Vic-Mallee)
Acid soil/aluminum toxicity (sNSW,
NE, and SW-Vic)
Frost
Salinity/sodicity tolerance
Manganese deiciency tolerance
Waterlogging tolerance
Preharvest spouting tolerance
Blackpoint/kernel discoloration
tolerance
Screening tolerance
Winter cropping only
Net form net blotch
Spot form net blotch
Scald
Powdery mildew
Leaf rust
BYDV
Net form net blotch
Spot form net blotch
Scald
Powdery mildew
Leaf rust
Stem rust
Common root rot
Crown rot
Root lesion nematode—Pratylenchus
neglectus and P. sp.
Root lesion nematode—Pratylenchus
thornei and P. sp.
Drought/terminal drought
Alkaline soil/boron toxicity tolerance
(WA-Mallee)
Acid soil/aluminum toxicity
Frost
Salinity/sodicity tolerance
Waterlogging tolerance
Drought/terminal drought
Frost
Preharvest spouting tolerance
Blackpoint/kernel discoloration
tolerance
Screenings tolerance
Winter cropping only
Preharvest spouting tolerance
Blackpoint/kernel discoloration tolerance
Screening tolerance
and parts of Victoria required medium-quick
maturing genotypes. Data from Queensland and
northern New South Wales revealed that medium
to medium-slow maturing European-type germ-
F
Winter/summer crop rotations, double
cropping
plasm gave better grain yield and quality in
eastern environments, but that quicker maturity
was required in western areas (•• Poulsen, unpub52
lished data).
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Barley Breeding History, Progress, Objectives, and Technology
Malting and feed quality
Quality attributes include both physical and
biochemical grain characteristics. Sparrow and
Doolette (1975) suggested that early assessment
of malting quality was aimed at identifying lines
with plump and uniform grain, high malt
extract, and quick and even germination when
malted. That deinition remains true today, and
a host of individual parameters is recognized as
contributing to each of the above attributes.
In 1983, the MBIBTC was formed to provide
industry guidance to the breeding programs
(Armitt and Way 1990). It has provided Australian
barley breeders with guidelines to develop new
malting cultivars (MBIBTC 1998) and has developed a rating system for malting barleys, which
takes into account weighted factors for malt
extract, protein modiication, diastatic power
(DP), wort viscosity, and dimethyl sulide content
(Armitt and Way 1990). The relative activities of
speciic enzymes such as α-amylase, β-amylase,
and β-glucanase and cellular constituents such as
β-glucan and starch are also recognized as target
parameters for malting barley improvement programs. Grain dormancy is essential to protect
against preharvest sprouting caused by summer
53 storms (Poulsen et al. 1995). Sprouting interferes
with the malting and brewing processes by causing
uneven starch and cell wall degradation in the
grain. The southern and western regions are less
concerned about dormancy as those areas have
predominantly winter rainfall patterns; however,
the southern coast of Western Australia is prone
to preharvest sprouting problems and a degree of
dormancy is required.
While some quality speciications are required
for all malting barleys, there is now a dichotomy
in domestic and export market requirements due
to the different sources of carbohydrate adjuncts
used in the brewing process (MBIBTC 1998).
The domestic brewing industry requires moderate levels of DP; however, many Asian breweries
require high DP values.
It has gradually become recognized that speciic quality attributes can enhance livestock performance. Johnston (1974) commented that barley
was seen as a source of energy and protein for
193
both the ruminant and nonruminant feeding
industries. He concluded that high protein
requirements of the feed industry could be better
dealt with through agronomic management than
breeding. Recent research has begun to deine
attributes of barley affecting animal feeding performance. Bowman and Blake (1996) identiied
considerable genetic variation in barley ruminant
feed quality. Dry matter digestibility, starch
digestibility, starch content, starch particle size,
ground grain particle size, and grain hardness
were identiied as key traits in cattle nutrition.
Resistance to biotic stresses
Early Australian priorities for disease resistance
breeding were scald (Rhynchosporium secalis),
covered smut (Ustilago segetum var. hordei), and
powdery mildew (Blumeria graminis f. sp. hordei)
(Sparrow and Doolette 1975). New South Wales
Agriculture produced scald-resistant lines from
the cross Commander × Prior during the 1940s.
However, named varieties were not released. In
the same era, covered smut and powdery mildew
resistant (Mlk; now susceptible) (from the Indian
variety Kwan) backcross-derived lines based on
Prior and Research were produced at the Waite
Institute (Pugsley and Vines 1946; Pugsley 1951),
leading to Prior A (=Kwan × 6* Prior; Kwan was 54
thought to have at least three genes conferring
resistance to covered smut), which has proven to
be a cornerstone of barley adaptation and improvement in southern Australia. Additionally, Prior
was one of the earliest of the European introductions and contains the Eam1 gene, which is prevalent in Australian germplasm (•• Franckowiak,
pers. comm.) and is the source of earliness in 55
Erbet (=Prior × 7* Betzes). This inherent “earliness” enabled grain ill to be completed in a hot
dry spring.
In the 1970s and early 1980s, disease resistance
was considered less important than grain yield or
grain and malting quality. Most programs used
disease-resistant parents, but effective resistances
were seen as desirable, rather than essential,
characteristics in new varieties (Johnston 1985;
Gilmour 1993). Disease resistance breeding took
higher priority in the New South Wales breeding
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program. Resistance to leaf diseases was seen as
necessary for early-sown cultivars grown in the
southern parts of the State (Read 1983).
Development of scald-resistant material was also
occurring in Western Australia (Portman 1983,
1985), South Australia (Sparrow 1983), and
Victoria (Ellis 1983).
By 1985, diseases were recognized as restricting barley production throughout Australia.
Hirsch (1985) estimated yield losses from diseases
in southern Australia between 1973 and 1983 to
be more than 16% of the overall crop value, with
a net cost of A$20–A$25 million. Rees (1985)
noted that considerable changes had occurred in
Queensland, and several diseases had become signiicant, including stem rust (Puccinia graminis),
leaf rust (Puccinia hordei), scald, net-type net
blotch (Pyrenophora teres f. sp. teres), spot-type
net blotch (P. teres f. sp. maculata), crown rot
(Fusarium pseudograminearum), and common root
rot (Cochliobolus sativus). This was in direct contrast to an earlier statement that the only regular
problem in northern barley crops was powdery
mildew (Rees et al. 1981). The increasing disease
levels were believed to have been due to changes
in seasonal conditions, pathogen genetic diversity, cultivars, and agricultural practices.
Severe stem rust occurred in Queensland
barley crops during 1982–1984 when the highly
susceptible cv. Galleon entered production on the
Darling Downs (Dill-Macky 1992). Localized
epidemics occurred in 1982 and 1984; however, a
more general epidemic occurred in the Darling
Downs and Burnett regions during the 1983
season. Shortly afterwards, Rees (1985) suggested
that the predominance of a single cultivar in
Queensland, Grimmett, was making the State’s
crop generally vulnerable to disease, in the event
that new pathogens or pathotypes became established. This threat was conirmed in 1988, when
new leaf rust pathotypes caused a signiicant epidemic in Grimmett crops. A combination of cultivar susceptibility and suitable weather conditions
led to highly damaging epidemics of leaf blotches
in 1998 (Poulsen et al. 1999; Rees et al. 1999). The
changing disease pressures in the northern region
56 led to changes in the NBIP breeding objectives.
A stem rust research program was initiated after
the 1982–1984 epidemics (Johnston 1985), and
other expansions to the program occurred after
the 1988 leaf rust and 1998 net blotch epidemics
(Poulsen 2001). In each case, projects were developed to assess the impact of the diseases and to
identify useful sources of genetic resistance
(Cotterill et al. 1992; Dill-Macky 1992; Platz 57
2001).
The most signiicant nematode pest of barley
in Australia has been CCN (Heterodera avenae),
which occurs naturally in South Australia and
Victoria. The southern programs have bred for
CCN resistance for many years (Ellis 1983;
Sparrow 1983). The irst Australian CCNresistant barley cultivar was Galleon, a South
Australian feed variety with the CCN resistance
gene Ha4 (Karakousis et al. 2003c) introgressed
from a North African landrace, CIho-3576 (Gall
eon = [Clipper × Hiproly] × 3* WI-2231, where
WI-2231 = Proctor × CIho-3576). The second
resistant cultivar was Chebec, which contained
the CCN resistance gene Ha2 (Barr et al. 1998)
from Orge Martin (Chebec = [Orge Martin × 2*
Clipper] × Schooner). Barleys are generally tolerant toward CCN, but resistance varieties reduce
the number of CCN in the soil, which has a
signiicant and positive effect on subsequent
intolerant and susceptible wheat crops.
There are no major insect pests at present in
Australia that can be addressed by genetic tolerance or resistances. Some pests such as Russian
leaf aphid are regarded as “quarantine threats,”
and there are prebreeding programs to overcome
potential incursions.
Resistance to abiotic stresses
Breeding for abiotic stress has generally been
associated with soil characteristics. In the 1980s,
the Waite Institute breeding program was selecting for manganese-eficient cultivars to address
deiciency problems in some South Australian
soils (Sparrow 1983). Conversely, the New South
Wales program, now followed by the BBA-West
program, has been developing cultivars adapted
to aluminum and manganese toxicity for production on acid soils (Read 1985). Parts of Victoria,
South Australia, and Western Australia have
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alkaline and sodic soils of marine origin, and these
are associated with boron toxicity. The breeding
programs in those States have developed conventional and molecular screening methodologies
and have incorporated boron tolerance into elite
germplasm (Jefferies et al. 1997; Barr et al. 2000).
inTEGRATiOn Of TECHnOLOGiEs
in AUsTRALiA
Barley breeding methods in Australia
Many barley breeders use pedigree breeding
systems. Speciic parents are hybridized by controlled pollination, and individual lines are
selected from segregating material in the F2 and
subsequent generations (Anderson and Reinbergs
1985). Selection usually commences with simple
traits and progresses to multienvironment yield
trials. Pedigree breeding strategies form the core
component of most, if not all, of the Australian
barley improvement programs. Various modiications may be applied to pedigree breeding strate-
195
gies. Riggs et al. (1982) designed a pedigree
program in which F4 bulks were tested in ield
trials to bring yield evaluation forward by 1 year
and to improve resource use eficiency in their
program. The NBIP uses a variation of this technique in its cross-evaluation strategy (Johnston
1983; Poulsen et al. 1995b).
The Western Australian program uses several
breeding strategies that are utilized where appropriate (Fig. 8.3.2).
The basic “conventional” method is an F2
progeny method with a reselection phase at the
F5. Almost 50% of the stage 2 program is derived
from doubled haploids (DHs). Marker-assisted
selection (MAS) coupled with DH, single-seed
descent (modiied, SSDm) and male sterile facilitated recurrent selection (MSFRS) are seen as the
methods of choice for increasing the rate of
genetic gain and for reducing the time to release
of new varieties.
Backcross breeding aims to recover as much of
a desirable recurrent genotype as possible while
introgressing a new gene from a less-adapted
parent (Harlan et al. 1922, cited by Anderson and 58
fig. 8.3.2. ••. A, agronomic; Y, yield; Q, quality; D, disease.
129
130
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Reinbergs 1985). In South Australia and Victoria,
a backcross strategy utilizing molecular markers
has been used to introduce disease resistance
genes and boron tolerance into the malting cultivars Sloop and Gairdner without signiicantly
changing their malting attributes (Barr et al. 2000,
2001).
Mass selection strategies are relatively inexpensive methods of early-generation screening
(Anderson and Reinbergs 1985). Australian
programs using this technique have included
New South Wales (B. Read, pers. comm.) and
Queensland (Poulsen et al. 1997).
Recurrent selection is an effective breeding
strategy used in self-pollinating crops to improve
complex characteristics such as yield, grain
quality, and partial disease resistance (Stuthman
et al. 1996). It involves repeated cycles of intercrossing and selection in a restricted gene pool.
Carver and Bruns (1993) concluded a review
paper by stating that “A review of genetic gains
(∆G) reported for grain yield and related traits
since 1985 indicates that recurrent selection has
been equally, if not more, effective than traditional breeding methods.” While ∆G in traditional pedigree programs was estimated at 1% per
year, recurrent selection programs with at least
two completed cycles achieved ∆G of 3.4% per
annum. In barley, a modiication of the technique
has been used in conjunction with a system of
genetic male sterility to make signiicant yield
improvements in North American feed barleys
(Falk 1996, 2001).
DH production through either anther culture
(Clapham 1973, cited by Kasha and Reinbergs
1982) or isolated microspore culture (Kasha et al.
1992) is used in Australia. However, DH production from the microspore-based techniques is
more attractive because of improved recovery of
DH lines, dificulties in managing Hordeum bulbosum plants, and spontaneous chromosome doubling, eliminating the need to use colchicine
(Broughton and Poulsen 1997).
The principal reason breeding programs use
DH systems is to save time (Logue et al. 1993).
As DH lines are, by nature, homogeneous, the
need for several generations of self-pollination to
achieve the same ends is eliminated. It is antici-
F
pated that 2–4 years can be saved from cultivar
development cycles in the Australian spring
barley breeding programs through the use of DH
techniques. For example, the DH cv. Tantangara
was released in 8 years as opposed to the conventional breeding cycle of 10–12 years in the New
South Wales Agriculture barley program (B.J.
Read, pers. comm.).
Interspeciic and intersubspeciic hybrids have
been used on several occasions to bring new genes
into cultivated barley. Hordeum vulgare subsp.
spontaneum has been used as a source of disease
resistance genes in many breeding programs
(Feuerstein et al. 1990; Abbott et al. 1991, 1992;
Chicaiza et al. 1996). Translocations between H.
bulbosum and H. vulgare subsp. vulgare have also
been used as a source of new genetic variations for
barley breeding (Xu and Kasha 1992; Pickering
et al. 1995, 1998, 2000). The scald-resistant
variety Tantangara, bred by Barbara Read at
NSW-DPI Wagga Wagga, has a resistance gene
from a H. vulgare spontaneum source (CPI-71283), 59
which was developed by Dr. Tony Brown (Brown
et al. 1993, 2000) as a Clipper backcross (BC3)
line, AB6 (Brown et al. 1988).
Disease resistance research and
breeding in Australia
Australia has diverse barley cropping environments, and consequently, the occurrence of barley
diseases is also diverse (Boyd and Dube 1989;
Murray and Brennan 2001). The soil-borne diseases, crown rot and common root rot, are widespread throughout Australia and have become
recognized in recent years as a national problem
(Murray and Brennan 2001). Scald and CCN (H.
avenae Woll.) have been major pathological constraints to barley production in southern Australia
for many years, and resistance has been virtually
mandatory in any new cultivars for the region.
However, net blotch and other stubble-borne
diseases have become increasingly signiicant
throughout the entire continent in recent years
(Murray and Brennan 2001).
There are ive rust diseases capable of infecting
cultivated barley; leaf rust (P. hordei), stem rust
(P. graminis f. spp.), and barley grass yellow rust
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(BGYR) (Puccinia striiformis) are the major cereal
rust pathogens of barley in Australia. Barley
stripe rust (P. striiformis f. sp. hordei) and crown
rust (Puccinia coronata Corda) have not been
recorded in Australian barley crops. Stripe rust is
considered to be a signiicant quarantine threat to
the Australian industry and preemptive breeding
strategies have been developed.
The sexual stages of P. hordei have been
reported on “Star of Bethlehem” (Ornithogalum
umbellatum) growing on the Yorke Peninsula in
South Australia (Wallwork et al. 1992). The
apparent sexual recombination of virulence genes
was described in this work, following isolation of
single leaf rust pustules collected from the area
and differential testing on appropriate barley
genotypes. This potential for genetic recombination of virulence factors and the consequent
development of complex new pathotypes is likely
to have a signiicant impact on resistance breeding
and on the deployment of resistance genes.
The Australian National Cereal Rust Control
Program (NCRCP) conducts annual surveys of
rusts in wheat, barley, oats, and other cereals
(McIntosh et al. 1995). The information is used
to monitor the development of new pathotypes,
recommend withdrawal of susceptible cultivars,
and strategically deploy resistance genes for the
most effective control of the diseases.
Severe localized epidemics of barley leaf rust
have been reported in Australia (Dill-Macky et al.
1989) associated with a predominance of leaf rustsusceptible cultivars. In Australia, leaf rust is a
signiicant disease of barley in southern
Queensland, in northern New South Wales, and
in parts of South Australia, West Australia, and
Tasmania (Murray and Brennan 2001).
The sporadic nature of Australian leaf rust epidemics is due to climate variability and luctuation in the relative production areas of susceptible
cultivars. The most recent epidemics of barley
leaf rust in Queensland and in northern New
South Wales occurred in 1978, 1983, 1984, and
60 1988 (•• Rees, pers. comm.). The 1988 epidemic
resulted from the development of two new
pathotypes (210P+ and 253P−) with virulence to
the resistance gene in the predominant cultivar,
Grimmett (Cotterill et al. 1994), causing substan-
197
tial yield losses of up to 30% (Dill-Macky et al.
1989). Two new and potentially damaging pathotypes of leaf rust (4610P+ and 5610P+) with virulence on most major cultivars were irst identiied
in Tasmania during 1991 (Cotterill et al. 1992) 61
and in West Australia in 1997 associated with the
varieties Triumph and Franklin (Loughman
1998). Park and Williams (2000) reported detection of the pathotypes in the northern region
during 1999 and that they made up over 60% of
the barley leaf rust isolates collected from across
Australia in the annual rust survey of that season.
The pathotypes are virulent on most Australian
barley cultivars and have caused economic
damage to crops in Western Australia and South
Australia.
Until 1999, Rph12 was effective against all
pathotypes of leaf rust found in Queensland and
Northern New South Wales. The irst Australian
report of virulence to Rph12 was conirmed with
pustules isolated from a Tasmanian crop of
Franklin barley during 1991 (Cotterill et al. 1992).
The new pathotype spread to South Australia by
1993 (H. Wallwork, pers. comm.). In 1998, a different pathotype, also with Rph12 virulence, was
reported from Western Australia (Loughman
1998). The two pathotypes designated 4610P+
and 5610P+ by octal notation were identiied in
South Queensland during 1999 (G. Platz, pers.
comm.). Rph12, derived from European cv.
Triumph, is present in barley cv. Tallon (Cotterill
et al. 1994) and Lindwall (R.G. Rees, pers.
comm.). Both of these are major commercial cultivars grown in the Queensland and northern
New South Wales winter cropping areas, and
thus, the northern region barley crop is at greater
risk from the disease. The release of Gairdner and
later Baudin, derived from either “Franklin sib”
or Franklin and both of which carry the Rph12
gene, has further exacerbated the problem of
having a longer-season, higher-rainfall barley
crop with a high proportion of susceptibility.
Backcrossing has been one of the more widely
used resistance breeding strategies in barley. This
is because the technique is simple and has a high
probability of developing well-adapted cultivars.
Backcross lines of Prior and Research were developed by the Waite Institute breeding program
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during the 1940s and were used as parents
(Pugsley and Vines 1946; Pugsley 1951). Later,
Western Australian breeders used backcrossing to
introduce net blotch resistance genes from
unadapted material from Ethiopia, Manchuria,
and Turkey into locally adapted genotypes (Boyd
et al. 1981). CCN resistance is expressed as a
dominant trait, and a limited backcrossing
approach has been utilized in both South
Australian and Victorian programs producing
such varieties as Galleon, Chebec, Sloop-SA, and
Sloop-Vic.
Recurrent selection is another strategy that can
be employed in barley to develop disease-resistant
germplasm (Sharp 1985). However, as opposed to
backcrossing, recurrent selection techniques
increase general resistance levels through several
cycles of recombination and selection in which
genes of minor effect are accumulated. Major
genes must not be present for recurrent selection
to work effectively, as they mask the minor gene
effects and interfere with the selection process. In
barley, recurrent selection has been successfully
used to improve resistance to leaf rust and
powdery mildew (Parlevliet and van Ommeren
1988). The MSFRS system developed by Falk
(1996) is now used routinely in the BBA-West
program utilizing both genotypic and phenotypic
screening methods. Genetic male steriles were
irst used in composite crosses (CC XIV) to facilitate hybridization (Ramage 1987 loc. cit.). Ramage
gave Eslick the credit for coining the term “male
sterile facilitated recurrent selection” (Ramage
1975).
It would seem that the best breeding strategy
for long-term control of barley leaf rust would
combine most of the systems described in the
two previous sections because of the diversity
of leaf rust pathotypes and the potential for
sexual recombination of virulence genes in the
pathogen. Australian barley pathologists and
breeders have had to deal with the problem that
the alternate host for barley leaf rust, Star of
Bethlehem (O. umbellatum), is present on the
southern end of Yorke Peninsula in South
Australia (Wallwork et al. 1992) and is probably
responsible for outbreaks of leaf rust in the
region.
F
The nBMMP and AWCMMP
In Australia, researchers participating in the
Australian NBMMP (Langridge 1997; Langridge
and Barr 2003, “Better Barley Faster: the Role of
Marker Assisted Selection” loc. cit.) produced
maps for 10 major populations and conducted
marker discovery in 44 minor populations (Barr
et al. 2001).
The major populations were selected to represent Australian genotypes and key parent lines.
They were Alexis × Sloop (DH/RIL) (Barr et al.
2003a), Amagi Nijo × WI-2585 (Pallotta et al.
2003), Barque*2 × H. vulgare subsp. spontaneum
71284-45 (advanced backcross quantitative trait
locus [ABQTL]) (J. Eglinton, pers. comm.),
Chebec x Harrington (DH) (Langridge et al.
1995; Barr et al. 2003b), Clipper × Sahara
(Karakousis et al. 2003b), Galleon × Haruna Nijo
(DH) (Langridge et al. 1995; Karakousis et al.
2003c), Mundah × Keel, (Long et al. 2003),
Sloop × Halcyon (DH) (Read et al. 2003),
Tallon × Kaputar (DH) (Cakir et al. 2003b),
Tallon × Scarlett (DH) (D. Poulsen, unpublished
data), VB9104 × Dash (DH) (D. Moody, unpublished data), VB9524 × ND11231 (DH) (Emebiri
et al. 2003), and Baudin × AC-Metcalfe (DH) (R.
Lance, pers. comm.). Maps of Chebec ×
Harrington and Galleon × Haruna Nijo have
been published on the Graingenes Web site along
with a Clipper × Sahara map (http://wheat.
pw.usda.gov/ggpages/maps.shtml).
One of the NBMMP objectives was to utilize
and improve new technologies for Australian
barley molecular geneticists. Simple sequence
repeat (SSR) markers were introduced into the
Australian barley research groups (Ablett et al.
2003; Karakousis et al. 2003a), and their application improved the eficiency and utility of markers
in breeding programs.
The availability of a substantial number of
markers common across individual barley maps
enabled the alignment of markers and the creation
of consensus maps of the barley genome. Mapping
data from six populations were combined to
produce the irst consensus map of 587 markers
(Langridge et al. 1995). The majority of the
markers were obtained from the Steptoe × Morex,
62
63
64
65
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Igri × Franka, and Proctor × Nudinka maps but
were linked together by common markers used
to map three Australian populations: Clipper
× Sahara 3771, Haruna Nijo × Galleon, and
Chebec × Harrington. Overall, the consistency in
gene order across the six original maps was high.
Discrepancies only occurred where markers were
tightly linked and were subject to the random
error normal to mapping experiments. A second
consensus map was constructed by Qi et al. (1996)
from the Proctor × Nudinka, Igri × Franka,
Steptoe × Morex, and Harrington × TR306
maps. The most recent barley consensus maps
were constructed by Karakousis et al. (2001,
66 2003d) using RFLP, AFLP, and SSR markers.
Among the early successes of the NBMMP
were the mapping of genes and markers for resistance to CCN, a major soil-borne parasite of
barley in southern production areas. Kretschmer
et al. (1997) reported the association of a single
dominant CCN resistance gene, Ha2, with lanking RFLP markers. The gene mapped to chromosome 2H. Molecular analysis of CCN-resistant
Australian cultivars demonstrated that all possessed the Ha2 gene. A second gene for CCN
resistance, designated Ha4, was mapped to chromosome 5H in Australian cv. Galleon (Barr et al.
1998). RFLP markers for both genes have been
used for selection in the South Australian program
(Barr et al. 2001).
QTLs for genes associated with high malt
extract were identiied and validated in populations between Australian and overseas barleys
(Collins et al. 2003). QTLs controlling kernel discoloration were described by Li et al. (2003a).
This information has been invaluable to select for
a dificult to measure trait associated with weather
damage at grain maturation and prior to harvest.
Similarly, QTLs controlling preharvest sprouting
have been identiied and implemented in breeding programs (Li et al. 2003b).
Box et al. (1997) reported evaluation of
markers with putative linkage to the gene for
hulless grain (nud) on chromosome 7H, and
marker assays were ultimately developed (A.
Box, pers. comm.). Markers for the hulless trait
enable the more eficient selection of progeny as
heterozygotes from various crossing situations.
199
Further studies identiied QTLs affecting coleoptile length on 7H and 5H in Proctor × Nudinka
and Galleon × Haruna Nijo, respectively (Box
and Barr 2000).
Four QTLs inluencing response to boron toxicity were mapped in Clipper × Sahara 3771
(Jefferies et al. 1999). Boron toxicity is a major
problem for barley crops in low-rainfall areas in
southern Australia. Two QTLs, located on chromosomes 4H and 6H, were associated with
reduced boron uptake. The 4H QTLs also
appeared to affect root length, dry matter production, and symptom expression. Leaf symptom
expression was affected by another QTL on 2H,
and suppression of root growth by high boron
levels was inluenced by a QTL on 3H.
In another series of molecular studies of
nutrient–plant relationships, a region on chromosome 4HL was identiied as contributing to
aluminum/acid soil tolerance in several genotypes (Raman et al. 2001, 2002, 2003). The
Australian data matched a locus mapped elsewhere (Tang et al. 2000). Raman et al. (2001)
identiied closely linked SSR markers for the
trait, with the intent of developing an alternative
to biological screening assays.
Mel1, a gene affecting the eficiency of manganese uptake in barley, was mapped to the distal
portion of chromosome 4HS (Pallotta et al. 2000,
2003). BSA of F2 plants from the cross Amagi 67
Nijo × WI-2585 was used to identify several
closely linked RFLP markers for the trait, which
was inherited from Amagi Nijo. Assessment of 95
genotypes used as parents in the South Australian
breeding program indicated that two RFLP
markers could be used to distinguish the presence
of the Amagi Nijo locus in all cases.
Consensus mapping was used to identify 62
SSR markers linked by less than 10 cM to 14
traits of interest to most Australian barley breeding programs (Karakousis et al. 2000). The traits
included resistance to powdery mildew (mlo),
net blotch (Rpt4), barley yellow dwarf virus
(BYDV) (Yd2) and CCN (Ha4). The markers
were validated with populations from the
NBMMP and successfully predicted plant phenotypes. Further analysis with breeding lines
from the South Australian Barley Improvement
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Program demonstrated that allelic variation was
suficient for the majority of the markers to be
used for routine screening.
68
Coventry et al. (2001, 2003a,b) examined
QTLs associated with yield and yield components
in the Australian mapping populations Alexis ×
Sloop, Chebec × Harrington, and Mundah ×
Keel. Several yield and yield component QTLs
were coincident with chromosomal regions known
to be associated with phenology and development, including the 2H Ppd-H1 photoperiod
response gene and another locus on the same
chromosome. The denso gene is present in the
Alexis × Sloop population and was associated
with major effects on grain size, yield, and plant
height. The phenological development patterns
of Australian barleys are an important aspect of
adaptation to Mediterranean and East Coast environments. Boyd et al. (2003) enhanced the classical understanding by investigating the molecular
genetic control of the basic vegetative period or
earliness per se and photoperiod response as they
apply to adaptation in the Australian context.
Apart from the loci discussed above, the
NBMMP mapped and developed markers for a
signiicant number of agronomic, quality, and
disease resistance characteristics. A mixed model
approach to better improve the QTL analyses
from multienvironment trials was investigated by
biometricians (Verbyla et al. 2003). Quality
research with respect to molecular approaches
was reviewed by Fox et al. (2003). Furthermore,
biometrics analyses of quality within breeding
programs and through quality laboratories from a
strategic point of view were studied by Cullis
69 et al. (2003). Other traits targeted by minor populations included grain staining, acid soil tolerance,
preharvest sprouting, boron tolerance, grain size,
and water sensitivity. Additional work conducted
as part of the NBMMP has been the validation of
markers for routine use in Australian breeding
programs. For example, Collins et al. (2001)
reported the validation of QTLs associated with
the expression of malt extract, DP, α-amylase,
and β-amylase in six representative breeding populations from the South Australian breeding
program. The work clearly demonstrated that
QTLs on chromosomes 1H, 2H, and 5H could be
used to improve malting quality through the
implementation of MAS.
Mapping barley disease resistance genes
in Australia
As a demonstration of the scope of the NBMMP
activities, disease resistance loci targeted in the
major mapping populations are described by
Williams (2003) and compared with other known
genes for resistance. In a more recent phase of
barley improvement in Australia, signiicant
attention has been placed on the development of
elite malting and feed barleys with multiple
disease resistances pyramided from complex
crosses.
Minor mapping populations were also constructed by the NBMMP and were analyzed by
BSA to identify markers linked to resistance to
scald (Genger et al. 2003), powdery mildew (Paris
et al. 2003), spot-type net blotch (Williams et al.
2003), net-type net blotch (Cakir et al., 2003; 70
Gupta et al. 2003; Raman et al. 2003), common
root rot, stem rust, leaf rust (Park et al. 2003),
crown rot, covered smut, spot blotch, and the
exotic quarantine diseases, such as stripe or yellow
rust (Cakir et al. 2003c).
Scald (R. secalis) has also been a signiicant
target for marker development as both the
pathotype structure and the genetics of resistance
are complex, making conventional selection techniques dificult. Rh, a gene conferring resistance
to scald, was mapped to chromosome 3HL by
BSA with RAPD markers in a set of near-isogenic 71
lines (Barua et al. 1993). It has been suggested
that the 3HL locus, now designated Rrs1, is multiallelic as is scald resistance in several other unrelated genotypes and has been mapped to the same
location (Williams et al. 2001). One of the alleles
confers resistance to 22 of the 23 known Australian
R. secalis pathotypes. The multiallelic nature of
Rrs1 has signiicant strategic implications in
designing crosses to combine scald resistance
genes.
Pathotype-speciic seedling resistance to nettype net blotch was mapped to chromosome 6H
in a DH population derived from the Australian
cross Tallon × Kaputar (Cakir et al. 2001, 2003a).
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The resistance was inherited from Kaputar, a
72 reselection of the CIMMYT cv. Arupo. Further
work is being carried out on the population to
identify the genetic location of adult plant resistance to net-type net blotch inherited from
Tallon.
A dominant resistance gene to spot-type net
blotch from Australian barley cv. Galleon was
mapped to chromosome 7H and was designated
Rpt4 (Williams et al. 1999). Flanking RFLP
markers were identiied for the locus. Validation
of the marker demonstrated greater than 90%
accuracy in predicting the occurrence of resistance to spot-type net blotch. Karakousis et al.
(2000) subsequently identiied SSR markers in
the vicinity of Rpt4, which have since been used
for selection in at least three Australian barley
breeding programs.
Other reports of marker development for
barley disease resistance genes include identiication of speciic markers and/or QTL for resistance to BYDV (Collins et al. 1996; Paltridge
et al. 1998). Stripe, leaf, and stem rust resistance
genes have all been targets for marker development and genetic study in barley. In general, the
development of markers has been seen as leading
toward the implementation of better breeding
strategies to develop long-term control of the diseases. The information obtained in these studies
has contributed to the development of a greater
depth of knowledge of the genetics and mechanisms of rust resistance.
Resistance to South American stripe rust
pathotypes was mapped to chromosomes 2H and
5H in Australian cv. Tallon (Cakir et al. 2001,
2003c). The data were obtained from a ield
assessment of the NBMMP Tallon × Kaputar
DH population at CIMMYT. Stripe rust resistance was also mapped in the NBMMP
Arapiles × Franklin mapping population, with
the resistance derived from Franklin (Cakir et al.
2003c). The Franklin resistance was also located
on 2H and 5H. The data suggested that the two
regions were common to both cultivars. It was
suspected that both Tallon and Franklin had
inherited the same resistance gene from Triumph,
situated on chromosome 5H. The location of the
QTL was not a perfect match for the 5H resis-
201
tance mapped by Chen et al. (1994), and further
study was required to determine the relationship
between the loci. The resistance QTL on 2H
from Tallon and Franklin appeared to be located
in the vicinity of a gene related to plant maturity
(Cakir et al. 2003). It therefore appeared that differences in maturity may have inluenced the
expression of resistance in the mapping populations at the time of observation.
Several leaf rust resistance genes from the Rph
series have been mapped and linked to molecular
markers including Rph2, Rph9/12, and Rph16
(Borovkova et al. 1997a,b, 1998; Graner et al.
2000). Poulsen et al. (1995a) identiied an RAPD
marker linked to the RphQ resistance gene from
barley accession Q21861. RFLP and STS markers 73
were subsequently used by Borovkova et al.
(1997a,b) to map RphQ to the centromeric region
of chromosome 5H, in the same region as Rph2,
and it was postulated on the basis of marker and
phenotypic data that Rph2 and RphQ were allelic.
Nine RAPD markers, two RFLP markers,
CDO749 and Rrn2, and one STS marker, ITS1,
were identiied as closely linked to RphQ
74
(Borovkova et al. 1997).
Molecular genetic marker
implementation in Australian barley
breeding programs
One of the major dificulties encountered in the
implementation of marker-assisted breeding in
Australian barley breeding programs has been the
initial availability of only a limited number of
suitable markers. Although RFLP markers were
used fairly extensively for CCN resistance and
boron tolerance by the Waite Institute program
in the late 1990s (A. Barr, pers. comm), other
Australian programs have taken longer to routinely adopt the technology. This has been partly
due to time requirements of developing and validating suitable PCR markers. Most Australian 75
programs are not fully equipped for the largescale operation of RFLP systems. However, with
the maturation of the NBMMP, the situation is
changing and all of the programs are reaching the
stage where routine marker implementation can
occur.
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The use of markers in Australian barley breeding programs was succinctly described in two
review papers (Barr et al. 2000, 2001). The majority of those markers were developed and/or validated within the NBMMP strategic initiative.
It was also reported that MAS had been used
in Australian breeding programs to select for malt
extract QTL on chromosomes 1H, 2H, and 5H
(Barr et al. 2001). In a particularly elegant
example, the South Australian program had used
selection for these loci in conjunction with a
“defect elimination” backcross breeding strategy
to develop improved versions of the cultivars
Sloop (Barr et al. 2000) and Gairdner (Barr et al.
2001). Markers were used to assist recovery of the
recurrent parent genomes while tracking genes
for BYDV resistance (yd2), spot-type net blotch
resistance (Rpt4), and CCN resistance (Ha2).
After the single genes had been introgressed,
selected lines were intercrossed with the aim of
developing versions of the cultivars possessing all
of the target genes.
However, the highest rate of usage of MAS in
Australian barley breeding has been the enrichment of complex crosses by the South Australian
program (Barr et al. 2000). By selecting F1 plants
with markers linked to a range of speciic traits,
the program is making more eficient use of material derived from three- and four-way crosses.
Only progeny lines with desired combinations of
loci are selected for ield assessment. F1 plants
from complex crosses requiring additional crossing can also be identiied. The new malting variety
Flagship from the University of Adelaide
program, accredited in 2006, is a testimony to the
eficiency and expediency of utilizing this
approach.
Marker use for the selection of QTL is also
progressing in the Australian breeding programs.
Collins et al. (2000, 2001) validated QTL related
to malt extract, DP, α-amylase, and β-amylase
and recommended markers for routine use in the
southern region breeding programs. Validation of
the markers in the northern and western region
programs has also commenced.
It is anticipated that the adoption of marker
technology by the Australian barley breeding programs will rapidly increase over the next few
F
years, especially with increased identiication of
PCR-based markers linked to traits of interest to
the individual programs.
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NEAR EAST, NORTH AND EAST
AFRICA, AND LATIN AMERICA
Salvatore Ceccarelli, Stefania Grando, and
Flavio Capettini
nEAR EAsT
Barley is grown on about 8 million hectares in 16
different countries in the Near East. However,
98% of the area is in six countries, namely,
Turkey, Iran, Syria, Iraq, Afghanistan, and
F
Pakistan (http://faostat.fao.org/, searched in 109
2008). There is a large variability in the types of
barley grown in this area. Two-row types predominate in Syria, Turkey, the northern part of
Iraq, and the rainfed areas of Iran, while six-row
types predominate in Pakistan and Afghanistan
and in the irrigated areas of Iraq and Iran. In most
of the dry areas of Syria and Iraq, and in some of
the rainfed areas of Northern Iran, farmers have
a strong preference for black-seeded types. Most
of the barley grown in the areas is either winter
or facultative and with a strong photoperiod sensitivity. With the exception of the irrigated areas,
barley is predominantly grown in the semiarid
and arid areas where rainfall is too low and erratic
to grow wheat. In the Near East, the predominant
use of barley is as animal feed. However, it is also
used as food in Iraq and Iran.
syria
Formal agricultural research in Syria began in the
early 1940s with the establishment of experiment
farms at Deir Elhajar and Kharabo, close to
Damascus. After independence, in 1946, various
Ministry of Agriculture and Agrarian Reform
(MAAR) directorates (such as horticulture, forestry, animal resources, and plant protection)
began to conduct limited agricultural research.
In 1964, the Directorate of Agricultural Scientiic Research (DASR) was established, taking on 110
responsibility for research activities under
MAAR. In 1982, DASR formed a technical committee to develop new varieties of cereals, legumes,
and forage crops and to coordinate research activities with the Department of Agricultural and
Scientiic Research (DASR), the International
Center for Agricultural Research in the Dry Areas
(ICARDA), and the Arab Center for Studies of
the Arid Zones and Dry Lands (ACSAD). This
could be considered as the beginning of the barley
breeding program, with training programs conducted on crossing techniques and breeding
methods.
In 2002, the nine existing agricultural research
entities under MAAR were merged to form the
General Commission for Scientiic and
Agricultural Research (GCSAR), which is now
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responsible for all the breeding programs (wheat,
barley, chickpea, lentil, faba bean, cotton, etc.).
The breeding methods used vary from the
introduction of international nurseries from
international centers such as ICARDA and from
other countries and their direct release as cultivars (examples in barley are “Furat 1” and “Furat
2”released directly from ICARDA’s international
nurseries); the introduction of segregating populations from ICARDA; the execution of crosses
followed by classical pedigree and the induction
of mutations (wheat, barley, chickpea, and soya
beans). Plant breeding programs give special
importance to traits such as drought resistance,
earliness, disease resistance, and high yield. The
barley breeding work is conducted at the stations
and centers of GCSAR, and neither foreign companies nor the private sector conducts plant
breeding activities in Syria.
The inal stage of the breeding program is the
on-farm trials, which are used to eventually recommend varieties for release. As a result of breeding work since the early 1970s, seven barley
varieties have been released, but their rate of
adoption is very low partly because of their lack
of adaptation to the farmers’ real conditions and
partly because of the limited capacity of the
General Organization of Seed Multiplication
(GOSM), which is only able to produce 10% of
the barley seed required for planting, while the
corresponding igure in the case of wheat is 60%
(Al-Ahmad et al. 1999).
Jordan
Formal agricultural research in Jordan began in
the early 1950s with the creation of the irst agricultural research station in the Jordan Valley.
During that decade, various other research stations were established throughout the country.
Research was initially carried out by the technical
divisions of the Ministry of Agriculture, which
were transferred to the Department of Scientiic
111 Agricultural Research when it was created in
1958, and then in 1970, the department
was merged with the Ministry’s extension unit
to form the Department of Scientiic Research
and Agricultural Extension. The Ministry of
209
Agriculture was restructured in the mid-1980s,
resulting in the creation of the National Center
for Agricultural Research and Technology
Transfer (NCARTT). In 2007, NCARTT was
renamed as the National Center for Agricultural
Research and Extension (NCARE). The Center
developed over the years, in part through the
National Agricultural Development Project
(NADP), which was cofunded by the United
States Agency for International Development
(USAID) and the Government of Jordan.
Agricultural research within the higher education sector began in the early 1970s with the
establishment of the Faculty of Agriculture (1972)
and the Marine Science Station (1974) at the
University of Jordan (UoJ). Even if research in
barley breeding is occasionally conducted at the
university, NCARE has the main responsibility
for breeding and releasing varieties (Taimeh and
Sunna 1999).
The barley breeding program in Jordan is
entirely based on introduced germplasm mostly
from ICARDA and ACSAD. The material is
tested in two research stations for 4 years before
being tested in on-farm trials and eventually recommended for release. In addition to the local
landrace (two-row), the most popular improved
variety has been Rum (six-row). Recently, the
program has released three additional varieties,
namely, Athroh (six-row), Yarmouk (Esp/18084L//Harmal, two-row), and Muta’a (Roho/A.
Abiad/6250, two-row).
During the last 4 years, the barley breeding
program in Jordan has gradually been converted
into a participatory program following the methodology described in Ceccarelli and Grando
(2007).
iran
Agricultural research in Iran dates back to 1925
when the Razi Institute, the “father” of the
current Razi Serum and Vaccine Research and
Production Institute (RSVRI), began its research
activity in the district of Karaj, about 35 km west
of Tehran. In 1926, the irst agricultural college,
afiliated with the Ministry of Agriculture, was
founded in Karaj. Important changes occurred in
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the 1950s and 1960s. New colleges of agriculture
were established and institutes were created with
technical support from international agencies,
such as the United Nation’s Food and Agriculture
Organization (FAO). Among these, the Seed and
Plant Improvement Institute (SPII) was established in Karaj in 1959 to conduct research and
seed multiplication on the main crops (cereals, oil
crops, cotton, rice, horticultural crops, forages,
etc.). In 1991, the Dryland Agricultural Research
Institute (DARI) was established in Maragheh
(Azerbaijan province) to relieve SPII from its
heavy responsibilities and to implement extensive
research on some of the major commodities and
noncommodity items of economic importance. In
particular, DARI conducts basic research on
mechanisms of abiotic (cold and heat) tolerance
from the physiological, genetic, and agronomic
points of view, and on developing strategies
to overcome these stresses and to increase
productivity.
Barley breeding started in Iran around 1930
with the exploitation of local germplasm. This
was followed in the 1950s by the beginning of
hybridization of local germplasm with exotic
germplasm and the extensive use of the pedigree
method, bulking in F6 and multilocation testing
in about 22 stations throughout the country. The
inal testing is done in the Uniform Regional
Yield Trials, which are speciically targeted to the
warm, cold, and rainfed regions of the country.
112
After the establishment of DARI, SPII kept the
responsibility for barley breeding in the irrigated
areas of the country, while DARI focused on the
rainfed areas (Roozitalab 1996).
Both breeding programs (SPII and DARI) collaborate actively with ICARDA: they receive,
test, and use both international nurseries as well
as special nurseries speciically targeted to the
growing environments in Iran. About 80% of the
barley germplasm is either a direct introduction
from ICARDA or is derived from crosses with the
ICARDA germplasm.
iraq
In the 1920s, a Directorate General of Agriculture,
afiliated with the Ministry of Economics and
F
Transport, started agricultural research activities
and established the irst experimental stations at
Abu Ghraib, near Baghdad and at Neinevah, near
Mosul.
In the 1940s, agricultural research activities
were conducted by the Directorate General for
Agricultural Research and Extension (DGARE)
until in 1958 the Directorate General for
Agricultural Research and Projects (DGAREJ)
was established. During the 1970s, research activities were expanded as several specialized research
stations and centers were established including
the Iraqi Atomic Energy Commission (IAEC). In
1980, the State Board for Applied Agricultural
Research (SBAAR) was established, which in
1987 was renamed as the State Board for
Agricultural Research and Water Resources
(SBARWS). In 1990, SBARWS was terminated
and replaced by the State Board for Agricultural
113
Research (SBAR) and CWSR.
Barley breeding in Iraq used to be divided into
two subprograms: one addressing the irrigated
areas around Baghdad and based in Abu Ghraib,
and the second addressing the rainfed areas of the
North, which, from an agroecological viewpoint,
are very similar to the areas in Northeast Syria.
The irst subprogram, based on six-row types
mostly bred for dual purpose as cattle feed, fully
exploited the segregating populations received
from ICARDA and released successful cultivars
such as IPA 7, IPA 9, and IPA 265. The second
subprogram, although addressing an area where
farmers’ preferences are for two-row black-seeded
barley, attempted the introduction of six-row
types. The major success of the program was the
release of Rihane-03, a straight introduction from
ICARDA, which in the late 1990s was cultivated
on about 250,000 ha.
Currently, due to the war since 2003, Iraq is
conducting a limited amount of variety trials
using material introduced from ICARDA.
Yemen
Agricultural research in Yemen dates back to the
1940s during the rule of the British Colonial
Government. El-Kod Research Station was
established in 1955 about 50 km from Aden, and
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Seiyun Research Center was established in Wadi
Hadramout in 1972 to cover the mid-altitude
region of South Yemen. Research activities were
later developed in most parts of the country, especially through numerous projects supported by
114 UNDP/FAO and IDA. They were introduced in
North Yemen starting in 1970, later developing
into a central research station in Taiz in 1978. In
1980, the Ministry of Agriculture and Agrarian
Reform of South Yemen created the Department
of Research and Extension (DRE), based at Aden,
which was transformed in 1986 to the Directorate
of Research and Extension (Ministry of
Agriculture and Fisheries 1989). At the same
time, the Agricultural Research Authority (ARA)
was established in Dhamar in North Yemen for
conducting research and applied studies to
improve agricultural production. In 1990, after
the uniication of North and South Yemen, the
Agricultural Research and Extension Authority
(AREA) was formed by merging DRE and
ARA and their respective research centers and
stations.
As in the case of other countries in the region,
Yemen’s barley breeding program is based on
introductions from international centers, particularly ICARDA and ACSAD, and more recently
from the International Atomic Energy Agency
(IAEA). Most of the breeding work takes place at
the research station of Al Erra, just outside the
capital city Sana’a. Two varieties, introduced
through ICARDA nurseries, Beecher and Arivat,
were released in 1986. In 1999, ICARDA and
AREA conducted a 2-year project introducing
participatory plant breeding in Yemen: two barley
varieties were adopted by farmers at the end of
the project, and currently, AREA is using participatory methodology to evaluate barley germplasm
obtained by mutation breeding from IAEA.
nORTH AfRiCA
Cereal cultivation in North Africa dates back to
ancient times. It seems that it preceded the
Phoenician colonization, which took place about
the twelfth century BC. The founding of Utica at
the delta of the Médjerda valley, particularly
211
suited to wheat cultivation, had the only objective
of supplying food for the long sea route from
Tyrus to Guadalquivir. Carthage eventually
replaced Utica as the export harbor near the
present site of Tunis.
When the Roman Empire, after destroying
Carthage, spread over North Africa, cereal cultivation was already established in the plateau of
Setif (Algeria) and in Numidia, which was later
occupied by the Romans because of the soil fertility and the abundance of cereals they could
provide. “The soil of Africa,” wrote Pliny, the
Elder in the irst century of our era, “has been
given as gift from Ceres: oil and wine were almost
refused; all the glory of the country is in the
harvest.”
Today, barley is second only to wheat in the
ive North African countries where it is grown on
about 3.5 million hectares (average of 2002–2006)
of which about 60% is in Morocco (http://
faostat.fao.org/). It is predominantly grown in
the semiarid and arid areas where rainfall is too
low and erratic to grow wheat. These include the
Northwest coast of Egypt, where high humidity
allows growing a barley crop with about 100 mm
annual rainfall, and in some provinces of Western
Algeria. In the latter, barley is an established crop
because of high demand for livestock feed.
Contrary to the Near East, the majority of the
barley landraces and improved varieties grown in
North Africa are six-row with very few exceptions. The release of superior two-row cultivars
has not been followed by any signiicant adoption
because of religious reasons (two-row types are
associated with beer production).
As in the Near East, the main use of barley is
as animal feed. However, the use of barley as
human food is much more widespread than in the
Near East, and in fact, Morocco has the world’s
largest per capita human consumption of barley.
In recent times, the consumption of barley as
human food has increased also in Algeria and
Egypt (Grando and Gomez Macpherson 2005). 115
The history of barley breeding in North Africa
varies among the countries, but, as already
described in the case of the Near East, in all countries is much more recent than for wheat. As a
consequence, and for many years, barley and
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wheat breeding were done by the same scientist,
in the same research stations, and with the same
philosophies and methodologies. This situation
did not consider that the two crops are grown in
distinct agroecologies by farms differing in size,
by farmers differing in wealth with access to differing amounts of information.
Tunisia
F
The history of barley breeding in Tunisia is
emblematic of most of North Africa and the Near
East. In Tunisia, cereal breeding started with
wheat at the beginning of the last century in the
then Service Botanique et Agronomique de
Tunisie (SBAT). Wheat breeding evolved from
the utilization of the variability existing within
local varieties of durum wheat, followed by mass
selection until 1930, when the irst crosses were
made. It was about at this time that bread wheat
was introduced from France and Algeria. Later in
the 1970s, the introduction of germplasm from
Centro Internacional de Mejoramiento de Maíz y
Trigo (CIMMYT) occurred, when several varieties were released, both using direct introduction
and selection from segregating populations. The
introduction of these varieties was accompanied
by the wider use of inputs such as fertilizers and
chemical weed control.
The barley breeding programs are much more
recent as the local varieties were used until 1950
when “Martin” and “Cérès” were introduced in
the North of the country. Problems with seed
production of these two varieties did not allow
replacing the local germplasm in the center and
the south of the country.
The irst serious attempt to start a barley
breeding program was made in 1973 (•• Harrabi,
116 pers. comm.) with crosses and selection of early
material for the semiarid environments. The
materials obtained were tested in central Tunisia
and in the high plateau of the Northwest. With
the establishment of ICARDA in 1976, Tunisia,
where ICARDA had a regional ofice, was used as
a testing site for ICARDA’s barley breeding
program, thus receiving more genetic material
than other countries. A large number of lines
were screened, and already in 1984, the irst
promising lines, such as ER/Apm, Roho, and WI
2198 (from the Waite Institute in Adelaide, South
Australia) were identiied and eventually released
in 1985 with the names of Faiz, Roho, and Taj,
respectively. The collaboration with ICARDA
has continued over the years with the exchange of
germplasm (both inished lines and segregating
populations) and the release of varieties based on
those nurseries.
Algeria
Wheat (bread and durum) and barley are the main
winter cereals in Algeria. Among the wheats, it is
possible to ind small amounts of Triticum turgidum, Triticum dicoccoides, and Triticum polonicum
in the local landraces of wheat and Triticum spelta
particularly in the oases.
As in Tunisia, farmers have initially grown
local landraces. Cereal breeding started in 1905
by Prof. L. Ducellier, who produced the irst pure
lines of durum wheat. After his premature death,
his work was continued by his students at the
Station Centrale d’Essais de Semences et
d’Amélioration des Plantes de Grande Culture de
Maison-Carrée and by other botanists as Trabut,
Bœuf, and Laumont.
The history of breeding can be divided into
117
three periods:
1.
2.
The irst period (1900–1962) was characterized by the utilization of the existing material
within local landraces with collections and
characterization of durum wheat. Laumont
and Ducellier started from 1930 the production of varieties selected by mass selection
procedure and then the irst crosses were
made; these crosses aimed at higher productivity and better quality.
The second period (1962–1990) was characterized by the introduction of germplasm
from Spain, Italy, Greece, Syria, Russia, and
the United States through European projects, FAO, CIMMYT, and then ICARDA.
Several varieties were released either by
direct introductions or selection from segregating populations. These varieties were
characterized by the need for more inputs
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(fertilizers and weed control) than landraces.
During this second stage, a limited crossing
program also started.
This irst stage was followed by the introduction
of durum wheats and by hybridization between
the local landraces and the introduced varieties.
Already at that time, the Station Centrale de
Maison-Carrée could count on a network of a
dozen regional experiment stations corresponding to the main cereal regions of Algeria.
The improvement of bread wheat, which
started being grown in Algeria only in 1930, followed the same route as durum wheat.
Similarly, in the case of barley, the initial cultivation of mixtures introduced from the Near
East was followed by pedigree selection, which
led to the development of locally adapted cultivars such as Saida 183 and Tichedrette, the latter
particularly adapted to the mountainous areas.
The majority of cultivated barley has white
kernels; however, farmers remember cultivating
black types considered resistant to drought.
Malting barley did not become important in
Algeria because of its susceptibility to shattering
and because of the indifference of the local
industry.
As a result of the collaboration with ICARDA,
several varieties were released, but the two locally
adapted types are still the most widely grown.
At the moment, Algeria’s barley breeding
program is almost entirely based on ICARDA’s
germplasm and in the last 3 years has started
being organized in a participatory way (Ceccarelli
and Grando 2007).
Morocco
The beginning of barley breeding in Morocco
dates back to 1920 and was based on the
improvement of local landraces and on the introduction of two-row foreign varieties. Among
hundreds of cultivars from Europe, the United
States, and Australia, about a dozen were selected
including Chevalier, Hannchen, Combesse,
Guldkorn, Princesse, and Prior (Grillot 1939).
Selection within the local populations initially
identiied two lines, 077 and 071, later named
213
Rabat 77 and Merzaga 71, respectively (Saidi
et al. 2005).
The introduction of two- and six-row varieties
characterized barley breeding until 1970 and led
to the release of cultivars such as Arig 8, Tamelalt,
Asni, and Azilal. Starting from 1980, the introduction of collections of barley from the United
States was the beginning of another phase of
barley breeding with the objective of improving
earliness and harvest index. The germplasm generated in this period was used for multilocation
testing to identify widely adapted lines (Amri
1993). Selection was conducted in the research
stations for disease resistance and yield. A dozen
varieties were released, but despite their performance, they were grown on no more than 5% of
the barley growing area. The poor adaptation to
farmers’ conditions was considered to be the
cause of the lack of adoption (Saade 1994). This
imposed a change of strategy, which consists of
breeding for wide adaptation in the case of favorable environments and for speciic adaptation in
the case of marginal environments. In the latter,
the barley breeding program in Morocco is now
also using participatory plant breeding.
Libya
Agricultural research in Libya dates back to the
early part of the twentieth century during the
Italian colonial era, when the “Centro Sperimentale
Agrario e Zootecnico della Libia” at Sidi El Masri
near Tripoli was established to serve the Italian
agricultural settlers.
In the early 1950s, the Ministry of Agriculture
was established and agricultural research became
afiliated with the Directorate of Plant and Animal
Production, with major changes in goals and
organization. In 1981, the General People’s
Committee (Minister’s Cabinet) initiated the
National Authority for Scientiic Research
(NASR) to formulate and supervise the national
research policy, to ill in gaps in research not
tackled by any existing research institutes and
centers, and technically to coordinate research
carried out at research centers. New research
centers were established under the umbrella of
NASR, including the Agricultural Research
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Center (ARC), where most of the plant breeding
is conducted. The Center has the following four
118 “Regional AR Centers” (RARC), supported by 13
experimental stations (Ben-Mahmoud 1966):
1.
2.
3.
4.
The RARC for the western area (which
encompasses the largest part of agriculture
and of the population) located at Tajura,
20 km east of Tripoli, has six research stations and three experimental sites and focuses
mainly on fully irrigated and supplementaryirrigated crops such as vegetables, cereals,
fruits, and forages, with some emphasis on
rainfed agriculture as well.
The RARC for the eastern area, located at
Al-Marj, 1100 km east of Tripoli, covers the
areas from Sirte to the Egyptian borders
(four stations)and focuses on rainfed agriculture (cereals, legumes, and fruits), forestry,
and range.
The RARC for the central area, located in
Musrata, 200 km east of Tripoli, covers the
areas from Khomes to Sirte (two stations). Its
10 researchers are working mainly on salinity
research.
The RARC located at Sebha (700 km south
of Tripoli) covers the southern area of the
country (one station) and works in the areas
of hot/dry climate and fully irrigated crops
(cereals, legumes, forages, and palm trees).
The breeding programs are mostly based on
testing germplasm introduced from international
organizations: in the case of barley, the main supplier of germplasm has been traditionally
ICARDA, which, as described below, in the last
10 years has begun distributing very early germplasm particularly suitable to the short growing
season of most areas where barley is grown in
Libya. Eight varieties have been released between
1992 and 2005.
Egypt
The irst school of agriculture in Egypt was established in 1869 and the irst directorate of agriculture in 1875.During the nineteenth century,
agricultural research was carried out by the
F
Egyptian Royal Society, and as early as 1897, a
number of experimental farms were established at
various locations. In 1910, the Agricultural
Authority was established with the responsibility
for conducting research and producing seed,
extending methods in crop production to farmers,
especially soil analysis and use of fertilizers, pest
control, and production of scientiic and technical
publications. The Ministry of Agriculture was
established in 1913. In 1957, research departments were formed, and one of the irst was the
Department of Plant Breeding (Rivera and Elkalla
1997).
The Ministry of Agriculture has undergone
several reforms in the past decades. It has grown
from only 7 departments in 1913 to 28 in 1950, to
194 in 1963, and to 92 these days dealing with
various aspects of agricultural production. Early 119
on among the major departments were Agriculture,
Horticulture, Plant Protection, Soil, Animal
Production, Veterinary Laboratories, and Seed
Production. These research departments were
reorganized in 1971 into one research body within
the Ministry of Agriculture and Land Reclamation
named the General Authority for Agricultural
Research, which was later (1983), renamed the
ARC, and evolved as the major institution for
agricultural research and extension in Egypt until
today and has the major responsibility for crop
breeding.
Barley breeding in Egypt has a very long
history, which started more than a century ago.
This has attracted the attention of international
organizations for the Egyptian genetic resources,
especially in relation to adaptation to abiotic
stresses such as drought, salinity, and poor soil
fertility. Many landraces and local varieties have
been collected from the desert areas of the
Northwest Coast, Siwa, and Sinai. These collections are maintained in gene banks and utilized by
different barley breeding programs for their valuable attributes.
Until the beginning of the last century, farmers
relied on their own local varieties developed by
selecting superior plants from existing landraces.
Each tribe had its own local variety (known as
Bedouin variety) selected and maintained by the
members of the tribe.
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Barley Breeding History, Progress, Objectives, and Technology
The irst organized efforts in barley breeding
started with the establishment of the Department
of Plant Breeding at Bahteem Agricultural
Research Station in 1898 as a part of the Sultanic
Agricultural Organization (Royal Agricultural
Society). Until the 1940s, barley breeding relied
mostly on the collection and selection from local
varieties and landraces known as “Baladi varieties.” These efforts resulted in the selection of
several high-yielding and disease-resistant varieties, such as Baladi 16, Bahteem 52, and Giza 24.
Starting in the 1940s, barley breeders started to
introduce germplasm from other countries, such
as Abyssinia 12, Hungaria 1, and Palestine.
Selection from this introduced germplasm
resulted in the release of two new varieties, Giza
68 and Giza 73, in 1948. In 1956, a new extremely
early barley variety (Hybrid 100) was developed,
which is also known as Saharawi 100. This variety
has been extensively used in the breeding program
for earliness until now. It was the irst barley
variety developed through crossing between
Baladi-16 and Atsel. Another important variety
was California Mariout, used side by side with
Sahrawi 100 in the rainfed areas of Egypt.
Barley breeders continued to cross local varieties and introductions to produce new varieties
with better adaptation to low rainfall areas. For
example, the variety Giza 117 was produced in
1959 as a selection from the cross Baladi
16 × Palestine 10, and the variety Giza 119 and
Giza 121, produced in 1973 and 1980, respectively, as selections from the cross Baladi
16 × Gem.
Another wave of barley varieties was developed
in 1980s, such as Giza 121, CC-89, Giza 123, and
Giza 124. The last two varieties replaced all the
old varieties because of their high yield potential,
especially in saline soils and under heat stress
conditions, respectively.
Barley breeding activities focus on screening of
barley genotypes for drought and salinity tolerance and for disease and aphid resistance as well
as heat tolerance. The germplasm is based on
introductions from ICARDA and on local breeding material developed from local crosses.
The program addresses three major environments, that is, the rainfed areas of the Northwest
215
Coast and Sinai, the new reclaimed land (irrigated) and saline soils, and the old irrigated areas.
The program follows a traditional pedigree
method with yield testing conducted on station in
micro and macro yield trials in several locations.
Seed multiplication is conducted at Sakha
research Station under irrigation.
In collaboration with the DRC, ICARDA has 120
conducted for the last 8 years a participatory
barley breeding program in the Northwest Coast:
this involves eight villages and has so far generated ive varieties, which are being multiplied by
farmers (by law, they cannot be oficially multiplied), and this is so far the main factor limiting
their wider adoption.
EAsT AfRiCA
The two most important countries for barley cultivation in East Africa are Ethiopia and Eritrea,
two of poorest countries in the world. In both
countries, agriculture is almost the only source of
living for the majority (85%) of the population.
Since Eritrea gained independence from Ethiopia
in 1991, the history of barley breeding was
common to the two countries before that date.
Barley is one of the most important staple food
crops and has a signiicant role in the diet of many
millions of people. Cultivation of slightly more
than 1 million hectares of barley occurs in the
highlands of the two countries. The farmers of
both countries grow barley because it has several
advantages over other cereals: (i) barley can be
grown in marginal areas where the choice of other
cereals is limited; (ii) it offers the farmer an earlier
crop harvest than most cereals, providing relief of
food shortages, which frequently occur during
the long rainy season; (iii) it has better stability of
production over other cereals; (iv) it is a dependable food crop as it is grown in different seasons
and production systems; (v) it is the preferred
crop for the preparation of traditional drinks and
beer; and (vi) its straw is a good source of feed
and bedding for animals and thatching of roofs.
Barley is believed to have been cultivated in
Ethiopia and Eritrea as early as 3000 BC. (Gamst
1969). Both countries, but Ethiopia in particular,
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have diverse climates, soils, topography, social
environments, vegetation cover, and livestock.
The long history of barley cultivation and the
diverse agroecological and cultural practices have
resulted in a large number of landraces and traditional agricultural practices. All the different
types of barley are grown: hulled, hulless, sixrow, two-row, irregular forms, dense, lax, hooded,
long awn, short awn, and rough and smooth awn
(Asfaw 1988). In addition, barley landraces vary
in their characteristics such as maturity, seed
color, seed size, seedling vigor, straw strength,
and in disease and insect pest resistance. Landraces
are also grown in mixtures, often with other
crops; the best known of these mixtures, called
hanfets, is a barley–wheat mixture very popular in
Tigray, the Northern part of Ethiopia, and in the
highlands of Eritrea (Woldeamlak et al. 2001,
2008).
Ethiopia and Eritrea are two of the centers of
diversity of barley, but also of other important
crops such as durum wheat, and therefore they
have a very long history of barley breeding if we
also include the millennia of selection conducted
by farmers.
Ethiopia
In Ethiopia, barley research started in the early
1950s by the former Alemaya College of
Agriculture and Mechanical Arts at its experimental station in Debre Zeit, with the evaluation
of landraces and introduced nurseries. After the
establishment of the Institute of Agricultural
Research (IAR) in 1966, barley research was
transferred from Debre Zeit to Holetta Research
Center.
After the restructuring of agricultural research
with the establishment of a federal center
(Ethiopian Agricultural Research Organization
[EARO]) and of agricultural regional centers,
barley research has been coordinated from the
Holetta Research Center in collaboration with the
regional research centers (at Kulumsa, Ambo,
Sheno, Adet, Sirinka, Mekele, and Sinana), and
other Ethiopian institutions. International collaborations include FAO, the United States
Department of Agriculture (USDA), and the
F
Swedish Agency for Research and Education
Cooperation (SAREC). Since the mid-1970s, the
collaboration with ICARDA has been valuable for
the exchange of information and germplasm and
for capacity building.
Between 1970 and 1990, nearly 14,000 local
landraces were evaluated. Most of the entries
were found susceptible to scald (caused by
Rhynchosporium secalis Oud.), net blotch (caused
by Helminthosporium teres Sacc.), spot blotch
(caused by Helminthosporium sativum Pum.), leaf
rust (caused by Puccinia hordei Otth.), and lodging.
However, from this work, some outstanding varieties such as Shege, Misratch, and Abay were
released in late 1990s (Lakew et al. 1997; Yitbarek
et al. 1998).
Between 1966 and 2001, nearly 30,000 exotic
entries were evaluated. Most of them were found
to be highly susceptible to scald, net and spot
blotch, and barley shoot ly (Delia lavibasis) and
had poor plant vigor and small grains. About 6%
were selected for further study. In the early 1970s,
the major contributors of germplasm were the
FAO Near East Regional Program, the USDA,
and the Arid Land Agriculture Development
(ALAD), and from the mid-1970s, ICARDA.
Germplasm has been also received from Brazil,
Colombia, the former Czechoslovakia, Egypt,
India, Kenya, Peru, Sweden, and the former
Republic of Yugoslavia (Gebre et al. 1996). From
these efforts, one hulled variety, AHOR 880/61,
was released, and some other elite lines are still
being used in the national crossing program as
sources of genes for desirable agronomic traits
such as grain quality and stiff straw and for disease
and insect pest resistance.
Between 1974 and 2001, over 1600 crosses
(single, double, and three-way) were made
between exotic and local germplasm and among
local germplasm to improve their resistance to
lodging and to major diseases such as scald, net
blotch, spot blotch, and leaf rust. F2 progenies
have been evaluated at a number of locations in
the country. Only one outstanding hulled barley
variety, HB 42, has been identiied from this
program.
Currently, some regional centers have started
programs of participatory barley breeding.
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Eritrea
In Eritrea, barley research started with a germplasm collection conducted by the staff of the
Department of Agricultural Research and Human
Resource Development (DARHRD) in 1997
(shortly after independence) and with the repatriation of local germplasm held in the Ethiopian
Gene Bank. Most of this work was funded by the
government of Denmark.
The barley breeding program started formally
in 1998 with the evaluation of the landraces collected in 1997 and with targeted crosses made at
ICARDA. In 1999, the irst ield trials were conducted in three villages, and three potential varieties were identiied. After an interruption caused
by the war with Ethiopia, the barley breeding
program started again and, since 2004, was funded
by a project of the Challenge Program on Water
121 and Food of the CGIAR.
In this second phase, after the systematic
evaluation of all the landraces available in the
gene bank, and of a number of experimental
hanfets, the National Agricultural Research
Institute (NARI), the new denomination of
DARHRD, has started the ield testing of new
breeding material derived from crosses made at
ICARDA between local landraces and exotic
germplasm.
LATin AMERiCA
Barley was introduced in the Americas by
Columbus as early as his second trip to the New
World. Barley was irst planted in 1493 by the
Spaniards who stayed at Isabela, Puerto Rico and
from there it was introduced to Mexico and the
United States. It is hard to document the irst
barley and wheat crops in South America, but it
most probably was in the Fort of Sancti Spiritus
in 1527, in the present day province of Santa Fe
in Argentina (Arias 1995). From there, it spread
to the Andes and the countries that share that
mountain range. The irst scientiic works in
barley and wheat in South America were carried
out starting in 1912 in Uruguay by Drs. Alberto
Boerger and Enrique Klein, selecting pure lines
217
resistant to leaf rust (gene Pa7) from populations
being used by farmers.
Barley production areas in Latin America can
be divided in two main groups, based on ecological regions and end use: (i) the Andean region,
with Bolivia, Ecuador, Colombia, and Peru,
where the main uses of barley are for food, feed,
and forage; and (ii) the countries of the Southern
Cone of South America, with Argentina, Brazil,
Chile, and Uruguay, plus México, where almost
exclusively barley is made into malt for beer production. The production in the irst group is
stable, whereas in the second, the production is
increasing due to the higher demand of malt
because of the rise in consumption of beer and
malt derivatives in the continent and worldwide.
After the establishment of the irst barley
research program in the subcontinent in La
Estanzuela, Uruguay, research has been carried
out at different times in different countries in the
region. Barley breeding at public institutions has
always been an appendix to wheat breeding and
has always counted on less economic and human
resources. Despite this, an important volume of
research and selection has been done in the Latin
American countries where barley is cultivated at
commercial scale (Arias 1995). At present, several
public and private long-term, well-established,
and successful breeding programs are working in
Argentina, Brazil, Chile, Ecuador, México, Peru,
and Uruguay. As the main varieties released were
for malting, very few local genetic resources have
been used in crossings because they were forage
and feed barleys. Almost all cultivated barleys
have spring habit, except in Chile, where some
winter or facultative barleys have been released.
Almost all cultivars are covered, except in Ecuador
and Peru, where a few hulless varieties have been
released.
In the Andean countries, epidemics of stripe
rust (caused by Puccinia striiformis f. sp. hordei)
introduced from Europe in the late 1970s affected
all varieties planted at that time, severely decreasing the production in all the highlands (Dubin
and Stubbs 1985). All the native cultivars were
highly susceptible, and the local breeding programs had to produce resistant cultivars in a relatively short period of time. At present, all released
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cultivars must show resistance to this disease in
the Andean countries and in Mexico.
ACKnOWLEDGMEnTs
The authors acknowledge the information on the
history of barley breeding in their respective
countries received from Dr. A.F. El-Sayed
(Egypt), Dr. M. Noaman (Egypt), Dr. S. Saidi
(Morocco), Mr. A. Shreidi (Libya), Dr. A.
Benbelkacem (Algeria), and Mr. M. Maatougui
(Algeria).
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Region. ICARDA, FAO, AARINENA, and CIHEAM,
Aleppo, Syria.
Amri, A. 1993. Comparison of performances of barley, wheat
and critical varieties. pp. 62–70. Project report IFAD/
ICARDA/Maghreb, Settat, January 1993.
Arias, G. 1995. Mejoramiento Genético Y Producción De
Cebada Cervecera En América Del Sur. Technical
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AUTHOR QUERY FORM
Dear Author
During the preparation of your manuscript for publication, the questions listed below have arisen.
Please attend to these matters and return this form with your proof.
Many thanks for your assistance.
Query
References
1.
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Query
Remarks
AUTHOR: Please conirm if all heading levels in this
chapter are correct.
AUTHOR: Please conirm if BaYMV should be deined
as “barley yellow mosaic virus.” Please conirm also if this
abbreviation is the same as BYMV. If so, please conirm
which of these abbreviations should be used in the
chapter.
AUTHOR: Fischbeck 1992 has not been included in the
reference list. Please supply full publication details.
AUTHOR: Barley Malt News 2009 has not been found in
the reference list. Please provide full reference details.
AUTHOR: Please check this Web site address and
conirm that it is correct. (Please note that it is the
responsibility of the author[s] to ensure that all URLs
given in this chapter are correct and usable.)
AUTHOR: Please conirm if the phrase “actual German
spring barley … superior malting quality” is correct.
AUTHOR: Please conirm if the deinition of QTL is
correct.
AUTHOR: Please deine CAPS.
AUTHOR: Please conirm f PCR should be deined as
“polymerase chain reaction.”
AUTHOR: Please deine GM.
AUTHOR: Please deine EMS.
AUTHOR: Please deine TILLING.
AUTHOR: Anonymous (2009): This entry has been
styled as a book chapter. Is this correct? If so, please
provide the chapter title and the editors’ surnames and
initials.
AUTHOR: Behn et al. 2004 has not been cited in the
text. Please indicate where it should be cited or delete it
from the reference list.
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AUTHOR: Friedt and Rasmussen. (2004): Please provide
the article title and the date of the proceeding (month,
day, and year).
AUTHOR: Friedt et al. (2000): This reference has been
styled as a book chapter. Is this correct? If so, please
provide the editors’ surnames and initials, the publisher
name, and the city of publication.
AUTHOR: Habekuss et al. (2009): Please check if the
abbreviated journal title is correct.
AUTHOR: Please provide the volume number and page
range for Humbroich et al. (2009) if available.
AUTHOR: Schmalenbach et al. (2009) has not been cited
in the text. Please indicate where it should be cited or
delete it from the reference list.
AUTHOR: Please provide the article title for Werner et
al. (2000).
AUTHOR: Werner et al. (2005) has not been cited in the
text. Please indicate where it should be cited or delete it
from the reference list.
AUTHOR: Werner et al. (2007) has not been cited in the
text. Please indicate where it should be cited or delete it
from the reference list.
AUTHOR: Please check the Web site addresses in this
chapter and conirm that all of them are correct. (Please
note that it is the responsibility of the author[s] to ensure
that all URLs given in this chapter are correct and
usable.)
AUTHOR: Poehlman 1978 has been changed to
Poehlman 1985 so that it matches the reference list. Please
conirm that this is correct.
AUTHOR: Please conirm if Septoria passerinii Sacc. Is
correct.
AUTHOR: Please conirm if the phrase “which gave
Canada wide importance for many years” is correct.
AUTHOR: Please deine CCRU.
AUTHOR: Roelfs and Huerta-Espino 1994 has been
changed to Roelfs and Huerto-Espino 1994 so that it
matches the reference list. Please conirm that this is
correct.
AUTHOR: Please check “to uptake suficient water.” Do
you mean “to take up suficient water”?
AUTHOR: Please conirm if the sentence “Finally,
nonmalting or feed barley … quality testing is not
required” is correct.
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31.
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AUTHOR: Please conirm if PCR should be deined as
“polymerase chain reaction.”
AUTHOR: Please conirm if the phrase “malt quality is
compared to the checked cultivars” is correct.
AUTHOR: Please check “The composition of these yield
trials … by a test coordinator.” Do you mean “The
composition of these yield trials, including checks, is
approved by the committee annually and the tests are
administered by a test coordinator”?
AUTHOR: Please provide the abbreviated journal title,
the volume number, and the page range for Chen (2004).
AUTHOR: Please update the publication year for Horsley
et al. (2008).
AUTHOR: Jorgensen (1992) has not been cited in the
text. Please indicate where it should be cited or delete it
from the reference list.
AUTHOR: Mornhinweg et al. (1999): Please conirm if
the page range is correct.
AUTHOR: Please provide the editors’ initials for Prom et
al. (1996).
AUTHOR: Weaver (1950) has not been cited in the text.
Please indicate where it should be cited or delete it from
the reference list.
AUTHOR: Sparrow and Doolette 1987 has not been
included in the reference list. Please supply full
publication details or delete these citations from the text.
AUTHOR: Please conirm if “The areas of greatest
production are … the Darling Downs of Queensland” is
correct.
AUTHOR” “mt” in all instances has been changed to
“MT.” Please conirm that this is correct.
AUTHOR: Please conirm if South Australia, Western
Australia, Victoria, New South Wales, Queensland, and
Tasmania should be abbreviated throughout the text as
SA, WA, Vic, NSW, Qld, and Tas, respectively.
AUTHOR: “Northern Region,” “Southern Region,” and
“Western Region,” respectively, have been lowercased
throughout the text. Please conirm if this is correct.
AUTHOR: AAB Grain 2008 has been changed to ABB
Grain 2008 so that it matches the reference list. Please
conirm that this is correct.
AUTHOR: Please conirm if the phrase “The result of
the signiicant investment … grain handling organizations
and authorities” is correct.
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AUTHOR: Please conirm if “Germplasm from the
NSWDPI, Victorian and Tasmanian breeding programs is
being progressed” is correct.
AUTHOR: Please deine HACCP.
AUTHOR: Please check the Web site addresses in the
chapter and conirm that they are correct. (Please note
that it is the responsibility of the author[s] to ensure that
all URLs given in this chapter are correct and usable.)
AUTHOR: Barr and Kneipp 1995 has not been included
in the reference list. Please supply full publication details.
AUTHOR: Please check if “Yield, lodging resistance, …
stem breakage” should be changed into a sentence.
AUTHOR: Poulsen, unpublished data: Please provide the
initial(s) of Poulsen.
AUTHOR: Poulsen et al. (1995): Please indicate whether
the year is 1995a or 1995b.
AUTHOR: Please conirm if “*” is correct or if it should
be changed to “×” in all instances.
AUTHOR: Franckowiak, pers. comm.: Please provide the
initial(s) of Franckowiak.
AUTHOR: Please deine NBIP.
AUTHOR: Dill Macky, 1992 has been changed to
Dill-Macky, 1992 so that it matches the reference list.
Please conirm that this is correct.
AUTHOR: Harlan et al. 1922 has not been included in
the reference list. Please supply full publication details.
AUTHOR: H. v. spontaneum has been changed to H.
vulgare spontaneum. Is this correct?
AUTHOR: Rees, pers. comm.: Please provide the
initial(s) of Rees.
AUTHOR: Cotterill, 1992b has been changed to Cotterill
et al. 1992 so that it matches the reference list. Please
conirm that this is correct.
AUTHOR: Please deine RIL.
AUTHOR: Please conirm that the full form of QTL is
correct.
AUTHOR: Please conirm if the deinition of SSR is
correct.
AUTHOR: Please conirm if the phrase “enabled the
alignment of markers and the creation of consensus maps
of the barley genome” is correct.
AUTHOR: Please deine RFLP and AFLP.
AUTHOR: Please deine BSA.
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AUTHOR: Coventry et al. (2003): “a” and “b” have been
inserted after the year of publication. Please conirm that
this is correct.
AUTHOR: Cullis et al. (2003) has not been included in
the reference list. Please supply full publication details.
AUTHOR: Cakir et al. 2003: Please indicate in all
instances in the text whether the years are 2003a, 2003b,
or 2003c.
AUTHOR: Please deine RAPD.
AUTHOR: Please deine CIMMYT.
AUTHOR: Please deine STS.
AUTHOR: Borovkova et al. 1997: Please indicate whether
the year is 1997a or 1997b.
AUTHOR: Please conirm if PCR should be deined as
“polymerase chain reaction.”
AUTHOR: Anderson and Reinbergs (1985): Please
conirm if the book title and the publisher names are
correct. Also, please provide the publisher location.
AUTHOR: Barley Australia (2008) has not been cited in
the text. Please indicate where it should be cited or delete
it from the reference list.
AUTHOR: Barley Breeding Australia (2008) has not been
cited in the text. Please indicate where it should be cited
or delete it from the reference list.
AUTHOR: Barr et al. (2001): Please conirm if the initial
of Barr is correct and please provide the month, day, and
year of the proceeding.
AUTHOR: Bowman and Blake (1996): Please conirm if
the book title and the publisher location are correct. Also,
please provide the publisher name.
AUTHOR: Box and Barr (2000): Please conirm if the
book title and the publisher location are correct. Also,
please provide the publisher name.
AUTHOR: Box et al. (1997): This reference entry has
been styled as a book chapter. Is this correct? If so, please
provide the editor name(s) and initial(s) and the publisher
name. Please conirm also if the book title and the
publisher location are correct.
AUTHOR: Boyd et al. (1981): This reference entry has
been styled as a book chapter. Is this correct? If so, please
provide the editor name(s) and initial(s), the publisher
name, and the publisher location.
AUTHOR: Broughton and Poulsen (1997): Please conirm
if the page range is correct.
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100.
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AUTHOR: Brown et al. (2000): Please provide the
publisher and its location.
AUTHOR: Cakir et al. (2001): Please provide the month,
day, and year of the proceeding.
AUTHOR: CBH Group (2008): Please conirm that this
reference entry is a book. If so, please provide the
publisher and its location.
AUTHOR: Chicaiza et al. (1996): Please conirm that this
is a book chapter. If so, please provide the publisher name
and please conirm that the book title and the publisher
location are correct.
AUTHOR: Coventry et al. (2003): Two reference entries
share the same author and year of publication. Hence “a”
and “b,” respectively, have been added to the years to
distinguish these entries in the text.
AUTHOR: Please provide the editor name(s) and
initial(s) for Dill-Macky et al. (1989).
AUTHOR: Emebiri et al. (2003): Please conirm that the
year of publication is correct.
AUTHOR: Falk (2001): Please conirm that this is a book
chapter. If so, please provide the publisher name and
please conirm that the book title and the publisher
location are correct.
AUTHOR: French and Schultz (1984a,b): Please conirm
if the article titles are correct.
AUTHOR: Graner et al. (2000): Please conirm if the
book title and the publisher location are correct and please
provide the publisher name.
AUTHOR: Jefferies et al. (1997): Please provide the
editor name(s) and initial(s), the publisher name, and the
publisher location.
AUTHOR: Johnston (1983): Please provide the editor
name(s) and initial(s), the publisher name, and the
publisher location.
AUTHOR: Karakousis et al. (2000): Please provide the
publisher name. Also, please conirm if the book title and
the publisher location are correct.
AUTHOR: Please provide the article title for Karakousis
et al. (2001).
AUTHOR: Kasha and Reinbergs (1982): Please provide
the publisher name. Also, please conirm if the book title
and the publisher location are correct.
AUTHOR: Kasha et al. (1992): Please provide the
publisher name and the publisher location.
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101.
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107.
108.
109.
110.
111.
112.
113.
114.
115.
116.
AUTHOR: Malting and Brewing Industry Barley
Technical Committee (MBIBTC) (1998): Please provide
the publisher location and please conirm if the publisher
name is correct.
AUTHOR: Poulsen et al. (1995b): Please provide the
publisher name and the publisher location.
AUTHOR: Poulsen et al. (1997): Please conirm if the
page range is correct.
AUTHOR: Poulsen et al. (2000) has not been cited in the
text. Please indicate where it should be cited or delete it
from the reference list. Also, please provide the publisher
name and conirm if the publisher location is correct.
AUTHOR: Raman et al. (2003): Please conirm if the
publication year is correct.
AUTHOR: Riggs et al. (1982): Please provide the
publisher name and the publisher location.
AUTHOR: Please provide the city location for Sharp
(1985).
AUTHOR: Stuthman et al. (1996): Please provide the
publisher name and please conirm if the publisher
location is correct.
AUTHOR: Please check this Web site address and
conirm that it is correct. (Please note that it is the
responsibility of the author[s] to ensure that all URLs
given in this chapter are correct and usable.)
AUTHOR: Please note that two deinitions of DASR
have been found in the text. Please conirm which of these
should be used in the chapter.
AUTHOR: Please conirm if “Department of Scientiic
Agricultural Research” is correct or if it should be
changed to “Department of Agricultural Scientiic
Research.”
AUTHOR: SPPI has been changed to SPII. Is this
correct?
AUTHOR: Please deine CWSR.
AUTHOR: Please deine UNDP and IDA.
AUTHOR: Grando and Gomez-Macpherson, 2005 has
been changed to Grando and Gomez Macpherson 2005 so
that it matches the reference list. Please conirm that this
is correct.
AUTHOR: Harrabi, pers. comm.: Please provide the
initial(s) of Harrabi.
F
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117.
118.
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123.
124.
125.
126.
127.
128.
129.
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131.
132.
133.
134.
135.
F
AUTHOR: Please conirm if “three periods” in the
phrase “The history of breeding can be divided into three
periods” should be changed to “two periods” as only two
periods were described in the text.
AUTHOR: Please spell out AR in “Regional AR
Centers.”
AUTHOR: Please check the phrase “and to 92 these days
dealing with various aspects of agricultural production.”
Should this be changed to “and to 92 at present, dealing
with various aspects of agricultural production”?
AUTHOR: Please deine DRC.
AUTHOR: Please deine CGIAR.
AUTHOR: Please provide the publisher location for Arias
(1995).
AUTHOR: Ben-Mahmoud (1966): Please provide the
place (city and country) where the workshop was held.
AUTHOR: Gebre et al. (1996): Please provide the
publisher and its location.
AUTHOR: Grando and Gomez Macpherson (2005):
Please provide the editor name(s) and initial(s).
AUTHOR: Please conirm that the abbreviated journal
title is correct in Rivera and Elkalla (1997).
AUTHOR: Please deine DUS.
AUTHOR: Fig. 8.1.4: “Author” has been changed to
“Source.” Is this okay?
AUTHOR: Please provide suitable legends for Figs. 8.3.1
and 8.3.2.
AUTHOR: Fig. 8.3.2: The deinitions of A, Y, Q, and D
have been deleted from the igure and moved to the igure
legend. Is this okay?
AUTHOR: Table 8.1.2: Please conirm if the area
harvested in Malta (400F) is correct.
AUTHOR: Table 8.1.2: Please conirm if the sentence
“Of the total global annual seed … (29%) in the EU (27)”
is correct.
AUTHOR: Table 8.1.3: n.d. has been deined as “not
detected.” Is this correct?
AUTHOR: Table 8.1.4: Please conirm if Pyrenophora
teres is correct.
AUTHOR: Table 8.2.2: The symbols †, ‡, and § have
been changed to superscript a, b, and c, respectively, as
per book style for table footnotes. Is this correct? Please
provide the meaning of these linking letters.
Ullrich—Production, Improvement, and Uses
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136.
137.
138.
139.
AUTHOR: Table 8.2.2: Please conirm if “db” should be
deined as “dry basis.”
AUTHOR: Table 8.3.1: Please conirm if the igure
“000 t” is correct.
AUTHOR: Table 8.3.2: Please conirm if the igure
“’000” in the table is correct.
AUTHOR: Table 8.3.4: Please spell out the genus name
in “P. sp.” in this table.
F
UBA 08
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