Principles of grafting and budding

農園系
黃建誌
The history of grafting

The origins of grafting can be traced to ancient times. There is evidence that the art of grafting was known to the
Chinese at least as early as 1560 BC.

Aristotle (384–322 BC) (亞里斯多德) and Theophrastus (371–287 BC) (泰奧弗拉斯托斯) discussed grafting in
their writings with considerable understanding.

During the days of the Roman Empire, grafting was very popular, and methods were precisely described in the
writings of that era. Paul the Apostle, in his Epistle to the Romans, discussed grafting between the “good” and the
“wild” olive trees (Romans 11:17–24).
Terminology (術語)

Grafting (嫁接) is the art of joining two pieces of living plant tissue together in such a manner that
they will unite and subsequently grow and develop as one composite plant.

接穗
嫁接接合區
砧木
Budding (芽接)
A form of grafting that uses a smaller scion piece— sometimes just a piece of the stem with
an axillary bud.

Scion (接穗) – piece of detached shoot with several buds and of the chosen variety that when attached to the
rootstock will become the shoot system of the grafted plant. The scion should be of the desired cultivar and free
from disease.
Rootstock (砧木)

Understock, stock
Lower portion of the graft that becomes the root system of the grafted plant; maybe seedling
or clonal

Interstock (intermediate stock, interstem)

piece of stem inserted by means of two graft unions between the scion and the
rootstock. Interstocks are used to avoid incompatibility between the rootstock and
scion, to produce special tree forms, to control dis- ease (e.g., fire-blight (火傷病)
resistance), or to take advantage of their growth-controlling properties.
Vascular cambium (維管形成層) is a thin tissue located between the bark (periderm, cortex, and phloem) and
the wood (xylem). Its cells are meristematic; that is, they are capable of dividing and forming new cells.

周皮

皮層 樹皮 Grafting terminology of the “bark” and

維管形成層
韌皮部 “wood” and associated tissues with
schematic drawing of a stem cross
section of a young woody plant stem.
後生木質部
髓
Callus (癒傷組織) is a term applied to the mass of parenchyma cells that develop from and around wounded
plant tissues. It occurs at the junction of a graft union, arising from the living cells of both the scion and
rootstock.
Rootstocks : seedling and clonal

Seedling rootstock

Seedling rootstocks propagated from seed can be mass-produced relatively simply and economically.
Viruses are transmitted from parent to progeny in very low percentages or not at all except in specific
instances. Seedling plants tend to have deeper rooted and more firmly anchored plants than rootstocks
grown from cuttings (e.g., plum and apple rootstock).

Seedling rootstock may show genetic variation leading to variability in growth and performance of the scion
variety. The variation can arise from natural heterozygosity of the source or from cross-pollination—both are
more likely if the rootstock is from an unknown, unselected source.
Rootstocks : seedling and clonal

Clonal rootstock
Clonal rootstocks are those vegetatively propagated by stool layering, rooted cuttings, or
micropropagation. Micropropagation of clonal rootstocks makes possible the production of great numbers of
such plants, upon which the scion cultivar can be grafted or budded.

Clonal rootstocks are desirable not only to produce uniformity, but also to utilize special characteristics such as
disease resistance. Clonal rootstock also influence the size and growth habit of the grafted plant and flowering
and fruit development of the scion.
An incompatible graft with the melon scion forming adventitious roots above the grafted Cucurbita rootstock.
Grafting and budding serve many different purposes

1. Perpetuating clones desired for their fruiting, flowering, or growth characteristics that
cannot be readily maintained or economically propagated by other asexual means.

2. Combining different cultivars into a composite plant as scion, rootstock, and interstock-
each part providing a special characteristic.

3. Changing cultivars of established plants (topworking), including combining more than one
scion cultivar on the same plant.

4. Repairing graftage for injuries-including inarching and bridge graftage.

5. Disease indexing—testing for virus diseases.

6. Study of plant developmental and physiological processes.

1. Perpetuating clones desired for their fruiting, flowering, or growth characteristics that
cannot be readily maintained or economically propagated by other asexual means.

-Cultivars of some groups of plants, including most fruit and nut species and many other woody plants,
such as selected cultivars of fir, eucalyptus, beech, oak, and spruce, are not propagated commercially by
cuttings because of poor rooting.

輻射松 南洋杉 濕地松 加勒比松 桉樹

2. Combining different cultivars into a composite plant as scion, rootstock, and interstock-each part providing a
special characteristic.

Obtaining the benefits of certain scions

Grafting selected cultivars can enhance plant growth rates, fruit characteristics and yield, and plant
form. “Weeping” forms of landscape plants can be obtained by grafting . Cactus and succulents
are easily grafted to produce unusual plant forms, as shown in

Weeping plant Cactus

2. Combining different cultivars into a composite plant as scion, rootstock, and interstock-each part providing a
special characteristic.

Obtaining the benefits of certain rootstocks

There are a number of benefits of grafting onto selected root- stock, including greater plant resistance
to biotic and abiotic stress, size control, enhanced reproductive growth, reduction in nursery
production time, and increased transplanting success.
Obtaining the benefits of certain rootstocks

Greater resistance to environmental stress and disease

Cleft-grafted-variegated English Holly on Ilex ‘Nellie Stevens’

rootstock adapted to the high temperature, periodic flooding,
low oxygen soils of the southeastern United States.
Obtaining the benefits of certain rootstocks

Greater resistance to environmental stress and disease

Pushing the ecological envelope. Using an inlay bark graft of

‘TifBlue’ blueberry (Vaccinium ashei) on a farkelberry
( Vaccinium arboreum ) rootstock, which tolerates a more basic
soil pH, allows the acid-loving blueberry to be produced in a
site with higher soil pH. (a) New scion growth with aluminum
foil and poly bag protecting the graft area. (b) Healed graft
union, and (c) ‘Tif Blue’ blueberry crop.
Obtaining the benefits of certain rootstocks

Greater resistance to environmental stress and disease

(a and c) Melon grafted onto boron-resistant

Cucurbita rootstock. (b) Non-grafted melon
showing boron susceptibility early in crop
cycle and (d) 1 month later.
Obtaining the benefits of certain rootstocks

Greater resistance to environmental stress and disease

Grafting vegetables is a common practice in Japan, Korea, the Mediterranean basin, and
Europe. It is used for managing soil-borne diseases, enhancing tolerance of low temperature and
salinity, and for increasing plant vigor and yield. (a) Grafted melon scion on curcurbita rootstock
with a grafting clip. (b) Melons grafted (white arrow) on Fusarium (鐮孢菌) -resistant
Curcurbita rootstock in Israel, (b) compared to susceptible, non-grafted melons (black arrows)
 Performance of Galia-type melons grafted on to Cucurbita 洋香瓜黑點根腐病
rootstock in Monosporascus cannonballus-infested and non-
infested soils

Wilt incidence of grafted and non-grafted

Galia-type melons in Monosporasus
cannonballus- infested soil
kg 10 m –2

Yields of grafted and non-grafted melons harvested in non-

infested soil.
(Cohen et al., 2005)
Grafting to manage soilborne diseases in Heirloom tomato production

青枯病(bacterial wilt)

2005

2006

(Rivard and Louws, 2008)

Controlling size of grafted plant

Relative size of apple trees on different rootstock

Controlling size of grafted plant
Hastening reproductive maturity
Hastening plant growth rate and reducing nursery production time

-In nursery production of shade trees, budded or grafted trees grow more rapidly than seedling or cutting-
produced trees; for example, Acer platanoides ‘Crimson King’ budded on a vigorous root-stock, and
budded Tilia cordata or budded Zelkova serrata grow more in 1 year than rooted cuttings will in 3 or 4
years

Zelkova serrata櫸樹
Tilia cordata小葉椴
Improving transplanting success

-Some plants rooted by cuttings make such poor root systems that they are difficult or impossible to
transplant; for example, the Koster spruce (Picea pungens) can be rooted in commercial numbers, but cannot
be successfully transplanted unless the root system is produced from grafted plants. Many Asiatic maples form
poor root systems from cuttings and must be grafted.

Picea pungens藍葉雲杉 Asiatic maples

 Obtaining the benefits of certain interstocks (Double-Working)

-Such a section is termed an interstock, interstem, intermediate stock, or intermediate stem

section. This is done by making two grafts, or double budding.

There are several reasons for using double-working in propagation:

1. The interstock makes it possible to avoid certain kinds of incompatibility.

2. The interstock may possess a particular characteristic (such as disease resistance or cold-hardiness) not possessed
by either the rootstock or the scion.
3. A certain scion cultivar may be required for disease resistance in cases where the interstock characteristics are the
chief consideration, such as in the control of leaf blight on rubber trees (Hevea).
4. The interstock may reduce vegetative growth and enhance reproductive growth of the tree. For example, when a
stem piece of the dwarfing ‘Malling 9’ apple rootstock is used as an interstock and inserted between a vigorous
rootstock and a vigorous scion cultivar, it reduces growth of the composite tree and stimulates flowering and
fruiting in comparison with a similar tree propagated without the interstock.
5. Obtaining special forms of plant growth. By grafting certain combinations together it is possible to pro- duce unusual
types of plant growth, such as “tree” roses or “weeping” cherry, birch, or willow cultivars.
Tree roses Weeping cherry
Changing cultivars of established plants (Topworking)

-A fruit tree, or an entire orchard, may be replaced with a more desirable cultivar. It could be unproductive, or an old
cultivar whose fruits are no longer in demand; it could be one with poor growth habits, or possibly one that is
susceptible to prevalent diseases or insects.

Topworking has sometimes been done by California producers of peach, plum, and nectarine every 2 to 3
years to take advantage of newer, more promising cultivars and thus remain competitive on the market.
TopWorking. (a) Inlay bark graft in top working an orchard. (b) Top worked citrus grove in Sicily using an
inlay bark graft. (c) Smaller citrus liner with inlay bark graft
印度棗主幹更新修剪嫁接

（邱祝櫻。2009）
Disease indexing—testing for virus diseases

-Virus diseases can be transmitted from plant to plant by grafting. This characteristic makes possible testing for
the presence of the virus in plants that may carry the pathogens but show few or no symptoms. By grafting scions
or buds on a plant suspected of carrying the virus onto an indicator plant known to be highly susceptible, and
which shows prominent symptoms, detection is easily accomplished. This procedure is known as indexing.

Sweet potato node with attached leaf, ready to be grafted on to an indicator plant for virus
indexing.

https://www.bioversityinternational.org/fileadmin/bioversity/publications/Web_version/117/ch5.htm
Thermotherapy

Thermotherapy is a heat treatment used to rid scion material of viruses. After the virus-free material is
indexed, as indicated previously, or tested with serological techniques, it can be multiplied by traditional
grafting/budding techniques.

(Wang et al., 2018)

Combination of thermotherapy and cryotherapy for the elimination of viruses that invade the
shoot meristem cells

(Bhojwani and Dantu, 2013)

In vitro micrografting of shoot tips of Citrus sinensis on a 2-week-old Troyer citrange
rootstock.

(Bhojwani and Dantu, 2013)

(Wang et al., 2018)
Study of plant developmental and physiological processes

-Grafting has enabled plant biologists to study unique physiological and developmental processes,
beginning in the 1700s with Stephen Hale’s studies on the circulation of plant sap.
Control of branching

Reciprocal grafts of rms-2 (K524) and WT (cv Torsdag)

plants. Arranged left to right are grafts of WT/WT (scion/stock),
WT/ rms-2, rms-2/WT, and rms-2/rms-2.

(Beveridge et al., 1994)

Adventitious root initiation

Grafting procedure and graft union

Root initiation in reciprocal grafts of H.

helix leaf cuttings after 33 d. From left to
right: mature/juvenile; juvenile/juvenile;
non-grafted juvenile; non-grafted mature;
mature/mature; juvenile/mature.

(Geneve et al., 1991)

Natural grafting

Occasionally, branches become naturally grafted together following a long period of being pressed
together without disturbance. In commercial orchards, limbs of fruit trees are sometimes deliberately
“braced” together and allowed to naturally graft, forming a stronger scaffold system to better support
the fruit load of the tree.

Natural grafting of roots is not as obvious but is more significant and widespread, particularly in stands
of forest species of pine, hemlock, oak, and Douglas-fir

pine Hemlock鐵杉
Natural grafting
-Fungal pathogens causing oak wilt and Dutch elm disease can be spread by such natural root
connections.

oak wilt 橡樹枯萎病 Dutch elm disease 荷蘭榆樹病

Axel Erlandson

https://en.wikipedia.org/wiki/Axel_Erlandson
Formation of the graft union

-Just as de novo meristems are necessary for adventitious bud and

root formation, a de novo-formed meristematic area (new vascular
cambium) must develop between the scion and rootstock if
successful graft union formation is to occur.
Formation of the graft union

-The development of a compatible graft is typically comprised of three major events

1. Adhesion of the rootstock and scion

2. Proliferation of callus cells at the graft interface or callus bridge
3. Vascular differentiation across the graft interface
1. Lining up vascular cambiums of the rootstock and scion

-The statement is often made that successful grafting requires that the cambium layers of rootstock and scion
must “match.” Although desirable, it is unlikely that complete matching of the two cambium layers occurs since
they are only one to several cell layers thick.

-In fact, it is only necessary that the cambial regions be close enough together so that the parenchyma cells
from both rootstock and scion produced in this region can become interlocked.
2. Wounding response

-A necrotic layer or isolation layer forms from the cell contents and cell walls of the cut
scion and rootstock cells.

-Undifferentiated callus tissue is produced from uninjured, rapidly dividing parenchyma

cells (薄壁細胞) (adjacent and internal to the necrotic layer). The callus tissue initially forms
a wound periderm.
3. Callus bridge formation
-Callus formation is a prerequisite for successful graft union formation. New parenchymatous callus
proliferates in 1 to 7 days from both the rootstock and scion.

Early callus bridge formation in Sitka spruce (Picea sitchensis) Sitka spruce錫特卡雲杉
Cross section of a Hibiscus wedge graft showing the importance of callus development in
the healing of a graft union

Hibiscus 木槿
4. Wound-repair xylem and phloem: differentiation of vascular cambium
across the callus bridge

-In both woody and herbaceous plants, the initial xylem and phloem are generally differentiated prior to the
bridging of vascular cambium across the callus bridge

Early vascular strands in callus bridge area

which are from wound-repair xylem and
wound-repair phloem.

Vascular connections after 14 days

5. Production of secondary xylem and phloem from the new vascular cambium in
the callus bridge

-The newly formed cambial layer in the callus bridge begins typical cambial activity, laying
down new secondary xylem toward the inside and phloem toward the outside.
Graft union formation in T- and chip budding

In T-budding, the bud piece usually consists of the “bark” (periderm, cortex, phloem, cambium), and often
some “wood” (xylem tissue). Attached externally to this is a lateral bud subtended, perhaps, by a leaf petiole. In
budding, this piece of tissue is laid against the exposed xylem and cam- bium of the rootstock.

Detailed studies of the grafting process in T-budding have been made for the rose , citrus, and apple.

T-budding https://youtu.be/_w051zyackM

chip budding https://www.youtube.com/watch?v=jGDgiJzmppY

More rapid union development in chip budding

In T-budding, the cambium of the rootstock is lifted in the flap of “bark,” so considerable callus in-filling and
development of new cambium must occur.
In part this advantage to chip budding is due to less callus filling being needed, and because there is no
requirement for an active cambium to lift the flap of rootstock bark, as there is with T-budding.

Detailed studies of the grafting process in chip budding have been made for the ‘Crimson King’ maple on
Acer platanoides (挪威楓) rootstock, ‘Conference’ pear (展會梨) on ‘Quince A’ rootstock, and ‘Rubra’ linden
on Tilia platyphyllos (寬葉椴樹) rootstock.

Acer platanoides (挪威楓) ‘Conference’ pear (展會梨) Tilia platyphyllos (寬葉椴樹)

Factors influencing graft union success

• Incompatibility
• Plant species and type of graft
• Environmental conditions during and following grafting
• Growth activity of the rootstock
• Polarity
• The craftsmanship of grafting
• Virus contamination, insects, and diseases
• Plant growth regulators and graft union formation
• Post-graftage—bud-forcing methods
Plant species and type of graft

-Some plants—including hickories (山核桃), oaks (橡樹), and beeches (山毛櫸)— are much more
difficult to graft than others even when no incompatibility is involved. Nevertheless, such plants,
once successfully grafted, grow very well with a perfect graft union.

-In grafting apples, grapes, and pears even the simplest techniques usually give a good
percentage of successful unions, but grafting certain stone fruits, such as peaches and apricots,
requires more care and attention to detail.
Successful saddle-
Unsuccessful
grafted grape with
saddle graft
profuse callusing
with rose
Environmental conditions during and following grafting
In apple grafts, little, if any, callus is formed below 0°C (32°F) or above about 40°C (104°F). At
32°C (90°F) and higher, callus production is retarded and cell injury increases with higher
temperatures. Cell death occurs around 40°C (104°F).
Environmental conditions during and following grafting

Moisture and plant water relations

-Air moisture levels below the saturation point inhibited callus formation; desiccation of cells
increased as the humidity dropped. In vitro studies of stem pieces of ash (Fraxinus excelsior)
have shown that callus production on the cut surfaces was markedly reduced as the water potential
decreased.

-With most plants, thorough waxing of the graft union or sealing of the graft union with polyethylene
grafting tape, Parafilm, or Buddy Tape helps retain the natural moisture of the tissues, which is all that is
necessary.
Fraxinus excelsior 歐洲白蠟樹 Buddy Tape
Growth activity of the rootstock
Some propagation methods, such as T-budding and bark grafting, depend on the bark “slipping,”
which means that the vascular cambium is actively dividing, producing young thin-walled cells
on each side of the cambium.

Chip budding can be done on a dormant or active rootstock.

Polarity in grafting
Polarity in grafting. In top-grafting , the proximal end of the scion is attached to the distal end
of the rootstock. In root grafting , however, the proximal end of the scion is joined to the
proximal end of the rootstock.

Inverse graft of grape with graft union

forming between the distal end of the
scion to the distal end of the rootstock.
Notice that the shoot reorients itself via
gravitational response.
Polarity in grafting

Bridge graft on a pear tree five months after grafting. Center scion was
inserted with reversed polarity. Although the scion is alive it has not
increased from its original size. The two scions on either side were
inserted with normal polarity and have grown rapidly.
Distribution-based morphogenic auxin activity gradients for embryo
patterning in plants

(Dhonukshe, 2009)
The craftsmanship of grafting

The art and craftsmanship in grafting and budding is critical for successful grafting. This is particularly
true with difficult-to-graft species, such as conifers (e.g., Picea pungens), which callus poorly, making
alignment of the cambial layers of the rootstock and scion critical. Conversely, the grafting technique is less
critical in grape or pear grafts, which callus profusely and have high grafting success

Picea pungens藍葉雲杉
Virus contamination, insects, and diseases
-Some delayed incompatibilities are caused by viruses and phytoplasma (mycoplasma-like organisms)

cherry leaf roll virus The cherry leaf roll virus causes blackline in walnut when it is initially
spread by virus-infected pollen of the symptomless English walnut
( Juglans regia波斯胡桃). The virus then travels down the scions of J. regia
into the susceptible rootstocks- California black walnut ( J. Hindsii 北
加州黑核桃) or Paradox walnut (J. hindsii × J. regia). The black walnut
rootstock (used for resistance to Phytophthora root-rot in the soil) has a
hypersensitive reaction and puts down a chemical barrier to wall-off the
virus, which causes the graft to fail, and a characteristic black line forms at
the graft union.
Blackline infection
Virus contamination, insects, and diseases

Apple union necrosis and decline (AUND) and brownline of prune is caused by the tomato
ring-spot virus that is transmitted by soil-borne nematodes to the rootstock and then to the graft
union.
Virus contamination, insects, and diseases

Top-grafting olives in California is seriously hindered in some years by attacks of the

American plum borer (Euzophera semifuneralis 美國梅螟), which feeds on the soft
callus tissue around the graft union, resulting in the death of the scion. In England,
nurseries are often plagued with the red bud borer (Thomasiniana oculiperda),
which feeds on the callus beneath the budshield in newly inserted T-buds, causing them to
die.

Euzophera semifuneralis 美國梅螟 Thomasiniana oculiperda

Plant growth regulators and graft union formation

Plant growth regulators, particularly auxin, applied to tree wounds or to graft unions give variable
results in wounding response and graft union formation . Auxin (IBA, NAA) and cytokinin (BA)
enhance graft success when applied to the base of side-grafted Picea scions, while the plant growth
retardant, dikegulac, stimulated scion growth by retarding rootstock development.
Post-graftage—bud-forcing methods
By “crippling” halfway through the rootstock shoot above the bud union and breaking the shoot over the
rootstock stem), girdling, or totally removing the rootstock above the scion bud union, apical dominance is
broken and the scion bud rapidly elongates.
偃枝栽培
Genetic limits of grafting

Grafting within a clone

A scion can be grafted back onto the plant from which it came, and a scion from a plant of a given
clone can be grafted onto any other plant of the same clone. For example, a scion taken from an
‘Elberta’ peach tree could be grafted successfully to any other ‘Elberta’ peach tree in the world.
Grafting between clones within a species

In tree fruit and nut crops, different clones within a species can almost always be grafted without
difficulty and produce satisfactory trees.

Douglas-fir (Pseudotsuga menziesii), incompatibility problems have arisen in grafting together

individuals of the same species, such as selected P. menziesii clones onto P. menziesii seedling
rootstock

Douglas-fir花旗松 Acer rubra 紅楓 Chinese chestnut 板栗

Grafting between species within a genus
For plants in different species but in the same genus, grafting is successful in some cases but unsuccessful in
others. Grafting between most species in the genus Citrus, for example, is successful and widely used
commercially.

But on the other hand, almond and apricot, both in the same genus, cannot be inter-grafted successfully. The
‘Beauty’ cultivar of Japanese plum (Prunus salicina) makes a good union when grafted on almond, but
another cultivar of P. salicina, ‘Santa Rosa,’ cannot be successfully grafted on almond.

Almond Apricot
Grafting between species within a genus

Reciprocal interspecies grafts are not always successful. For instance, ‘Marianna’ plum
(Prunus cerasifera × P. munsoniana) on peach (Prunus persica) roots makes an excellent graft
combination, but the reverse—grafts of the peach on ‘Marianna’ plum roots— either soon die or fail
to develop normally.

Prunus cerasifera × P. munsoniana Prunus persica

Grafting between genera within a family
When the plants to be grafted together are in the same family but in different genera, the chances of a
successful union become more remote.

However, high success rates occur between Nootka cypress (Chamaecyparis nootkatensis) grafted on
Chinese arborvitae (Platycladus orientalis) rootstock

Cupressaceae 柏科
Nootka cypress 阿拉斯加扁柏 Chinese arborvitae 扁柏
Grafting between genera within a family

The quince (Cydonia oblonga) has long been used as a dwarfing rootstock for certain pear (Pyrus communis and
P. pyrifolia) cultivars. The reverse combination, quince on pear, though, is unsuccessful. The evergreen loquat
(Eriobotrya japonica) can be grafted on deciduous and dwarfing quince rootstock (Cydonia oblonga)

Rosaceae 薔薇科

Cydonia oblonga 榲桲 Pyrus communis 西洋梨 Pyrus pyrifolia 沙梨 Eriobotrya japonica 枇杷

Grafting between families

Successful grafting between plants of different botanical families is usually considered to be

impossible, but there are reported instances in which it has been accomplished.

These are with short-lived, herbaceous plants, though, for which the time involved is relatively brief.
Grafts, with vascular connections between the scion and rootstock, were successfully made using white sweet
clover, Metilotus alba (Leguminosae) as the scion, and sunflower, Helianthus annuus (Compositae) as the rootstock.

Fabaceae 豆科 Asteraceae 菊科
Metilotus alba 白花草木樨 Helianthus annuus 向日葵

(Nickell, 1946)
Graft incompatibility
The ability of two different plants, grafted together, to produce a successful union and to develop
satisfactorily into one composite plant is termed graft compatibility.

Graft failure can be caused by anatomical mismatching, poor craftsmanship, adverse

environmental conditions, disease, and graft incompatibility.

Graft incompatibility occurs

(a) Adverse physiological responses between the grafting partners

(b) Virus or phytoplasma transmission
(c) Anatomical abnormalities of vascular tissue in the callus bridge
Graft incompatibility in ‘Jonagold’ apple scions budded to dwarfing ‘Mark’ rootstock

(a) Unstained section, with callus tissue between the rootstock and scion.
(b) Section stained with toluidine blue O (甲苯胺藍). The xylem (x) in the graft union is interrupted by
parenchyma tissue (薄壁組織) (arrows) which limits water flow and survival of the scion
Graft incompatibility Breakage at the graft union resulting from
incompatibility

(a) Compatible apple chip bud with vascular

continuity indicated by red dye, azosulfonate.
(b) Unsuccessful chip bud with vascular discontinuity,
as indicated by no visible dye (a) One-year-old nursery trees of apricot on almond
seedling rootstock.
(b) Fifteen-year-old ‘Texas’ almond tree on
seedling apricot rootstock, which broke off
cleanly at the graft union—a case of “delayed
incompatibility” symptoms
Graft incompatibility

Graft incompatibility occurring some 15-plus years after the Monterey pine (輻射松) ( Pinus radiata ) was
grafted.
External symptoms of incompatibility

• Failure to form a successful graft or bud union in a high percentage of cases.

• Yellowing foliage in the latter part of the growing season, followed by early defoliation.
Decline in vegetative growth, appearance of shoot dieback, and general ill health of the tree,
including drought stress
• Premature death of the trees, which may live for only a year or two in the nursery.
• Marked differences in growth rate or vigor of scion and rootstock.
• Differences between scion and rootstock in the time at which vegetative growth for the season
begins or ends.
• Overgrowths at, above, or below the graft union.
• Suckering of rootstock.
• Graft components breaking apart cleanly at the graft union.
External symptoms of incompatibility

Graft compatibility affects water uptake.

(a) Arava melon showing hotter scion and cooler temperature in Cucurbita rootstock (arrow) with
noncompatible grafting combination.
(b) Compatible graft showing uniform temperature between scion and rootstock. Differences in
temperature gradients determined with a thermal camera.
External symptoms of incompatibility

Physiological incompatibility between scion and rootstock. Scion

overgrowth caused by blockage of assimilates translocating from the
scion to the rootstock, causing a weak root system.
External symptoms of incompatibility

While rootstock outgrowth is not desirable, a large,

strong tree can still develop.
(a) Sweet orange rootstock used for
dwarfing, overgrowing the grapefruit scion.
(b) Rootstock overgrowing scion on Morus
alba ‘Platanifolia.’
External symptoms of incompatibility

Scion or rootstock outgrowth can still lead to a large, strong tree.

Such outgrowth (arrows) is more related to the genetic tendency for
growth, than to incompatibility.

(a) Scion overgrowing rootstock: Acer

pentaphyllum (五小葉槭) on A.
pseudoplatanus (大槭樹) rootstock, and (b)
grapefruit scion on sour orange rootstock,
which tolerates alkaline, heavy soils, but
can be susceptible to Trestiza
Suckering of rootstock

Undesirable suckering of rootstocks. (a) Hamamelis vernalis (北美金縷梅) ‘Sandra’ grafted on

Hamamelis vernalis rootstock, and (b) rootstock suckers on recently grafted Ulmus alata (翅榆)
‘Lace Parasol’ grafted onto seedling Ulmus alata . The suckers will need to be removed.
Pathogen-induced incompatibility
Viruses and phytoplasmas (mycoplasma-like organisms) cause pathogen-induced
incompatibility.

Latent viruses in the scion portion of graft combination may

cause symptoms to appear in a susceptible rootstock following
grafting.
Causes and mechanisms of incompatibility
Physiological and biochemical mechanisms

One proposed physiological and biochemical mechanism concerns incompatible combinations of certain pear
cultivars on quince rootstock. The incompatibility is caused by a cyanogenic glucoside (氰苷), prunasin, normally
found in quince but not in pear tissues.
Prunasin (洋李甙):含氰配醣體
Effects of the rootstock on the scion cultivar

-Size and growth habit

Fruiting: Fruiting precocity, fruit bud formation, fruit set, and yield of a tree can be influenced by the
root- stock used.
Effects of the rootstock on the scion cultivar
Size, quality, and maturity of fruit

Stock-scion relations.
(a) Seedless ‘Marsh White’ grapefruit scion on rough lemon
rootstock
(b) ‘Marsh White’ on sour orange rootstock, which has a
thinner peel (arrows)—and is also sweeter and juicer

rough lemon 粗皮檸檬 sour orange苦橙