J. J. SCHUURMAN AND M.A.J.

GOEDEWAAGEN

METHODS FOR THE

EXAMINATION OF
ROOT SYSTEMS
AND ROOTS
Methods in use at the Institute
for Soil Fertility for eco-morphological
root investigations

/<
Dr. J. J. Schuurman is a senior scientific officer at the Institute for Soil Fertility,
Haren (Gr.) the Netherlands.
The late Dr. M.A.J. Goedewaagen was a senior scientific officer at the same
institute until his retirement in 1957.
He died in 1970.

BIBLIOTHEEK
DER
LANDBOUWHOGESCMOefc
W*GKNINGEN

ISBN9022003248

©Centrefor AgriculturalPublishing and Documentation, Wageningen, 1971

No part of this book may be reproduced and/or published in any form, by print,
photoprint, microfilm or any other means without written permission from
the publishers.

Printed in the Netherlands by Drukkerij Van Dooren N.V., Vlaardingen

Contents

Preface 1

l. Introduction 2
l.i Morphological characteristics 2
1.2 Types of technique 3
1.2.1 Methods in field trials 3
1.2.2 Methods in container trials 4

2. Monoliths: the pinboard method 4

2.1 Equipment 4
2.2 Obtaining pinboard samples 5
2.3 Removing soil from the root systems 8
2.3.1 Sandy or loamy soils 8
2.3.2 Heavy clays 11
2.4 Final steps 12
2.5 Photography 13
2.5.1 Root systems , 13
2.5.2 Shoots 15
2.6 Calculation of root weights 16
2.7 Description of root systems 16
2.8 Description of individual roots 16
2.8.1 Selection of roots 16
2.8.2 Preparing roots for photography 17
2.8.3 Photography 19
2.8.4 Description 20
2.9 Special applications and difficulties 21

3. Soil specimens having a slight volume: the auger method 22

3.1 Equipment 22
3.2 Sampling 25
3.2.1 The light auger / 25
3.2.2 The heavy auger 27
3.3 Calculation of root weights 28
3.3.1 Hand washing 28
3.3.2 Routine method 32
3.4 Results 35
3.5 Field assessment ofroot densityin cores 36

3.6 Special applications and difficulties 39

4. Excavations 43

5. Profilewalls 44
5.1 Mappingfollowed bycounting 44
5.2 Weaver's method 49
6. Experimentsincontainers 49
6.1 Cylinders 50
6.1.1 Material 50
6.2 Largecylinders 51
6.2.1 Filling 51
6.2.2 Soilwatertable 54
6.2.3 Sampling , 54
6.3 Smallcylinders 59
6.3.1 FiHiwg 59
6.3.2 Soilwatertable 60
6.3.3 Sampling 61
6.4 Special applications 61
6.5 Casesandboxes 62
6.6 Pot cultures 64

6.7 Nutrient solutions 66

7. Tracers 67

Appendices 68

Bibliography 71

Keytobibliography 85
Preface

In 1911Maschhaupt initiated studies on root morphology at the Institute

for Soil Fertility at Groningen. Zijlstra worked briefly in this field but
discontinued these studies. Goedewaagen started work at the Institute in
1921and spent 41years there working on root morphology, publishing most
of his results in Dutch. The experience over these years led to considerable
development of morphological and ecological methods of research on roots,
whichwere compiled into two small publications by Goedewaagen in 1948,
1949.
Afuller account, the first edition of this book, was published in 1965 but
has sold out and the publisher has asked me to prepare a second revised
edition. This second edition includes new findings, in particular new
techniques since 1965.For auger soil cores, a new routine partly automatic
method of cleaning roots is described. The schemes for description of roots
have been transferred to appendices and the bibliography has been brought
up to date. There are more figures and plates. However, as modern materi-
als are not always at hand, old methods have usually been kept in the book.
Root ecology may stress physiology or morphology. This book is confined to
methods of morphological research.
Irealize that my account is sometimes incomplete and will be happy to
supply readers with further information. The methods described are as used
in our institute. Many details are adaptable to circumstances elsewhere and
to materials available. I am fully aware of the existence of other methods.
Since description of these would have unduly enlarged the size of this book,
an extensive literature list is given with akeythat can be consulted by
those who want more information.
Once again Ithank Mr.J.J. H. de Boer and Mr. L.Knot for helping to
prepare this edition. Mr. G.Mesker has produced many original ideas in the
development of the new method of cleaning roots in soil cores. MissA.C.
G. Holle is largely responsible for the revision of the bibliography. Mr. J.
Floris has helped to develop methods for examining finer details of roots.
Thanks are also due to Mrs.E.Brouns and Mr. J. C.Rigg'of Pudoc for
English correction.

Haren-Groningen, March 1971

J.J. Schuurman

1
1. Introduction

Plant growth is governed by different processes: synthesis of organic

material by green aerial parts, and uptake of nutrients and moisture by
roots. Theseprocesses interact in that carbon dioxide assimilation is
necessary for root growth and uptake by the roots for shoot growth.If
fixation of carbon dioxide is poor, roots grow badly.Reciprocally if roots
absorb only small amounts of nutrients and moisture, aerial growth is
impaired even when aerial conditions are favourable. Symptoms observed
above the ground may be caused by soil factors that affect the development
of the root system and consequently the absorptive capacity of the root
system.
Besides the content of nutrients and moisture in the soil and the vigourof
the root system, absorptive capacity is determined by the distribution and
extent of the root system. Agricultural crops usually need a well developed
root system in order to exploit deeper layers more fully. They are then
more resistant to periods of stress that often occur during growth and are
more likely to yield well.Farmers should know what factors promote and
impede root growth. With this knowledge, they can purposefully encourage
root growth. Such help to the farmer is the ultimate aim of ecological
research on roots. In root ecology, certain morphological characteristics are
important (Section 1.1). They can be studied in the field or in containers.
In either case there are problems in making roots accessible for study
without damaging them. Methods will be described in later chapters in the
same order as enumerated below (Section 1.2).

1.1. Morphological characteristics

1. Total amount of root. This indicates absorptive capacity. But two plants
are comparable onlyif they have similar types of root, e.g. plants of the
same species of perhaps the same family. The total amount of root is
usually expressed in grams dry root but can also be expressed as total
length of root or as total surface area. The total surface area is an impor-
tant characteristic, but it is extremely difficult to calculate.Anyway, not
everypart of the surface is functionally identical. It is therefore ignored.
Total weight of root or root weight in the surface soil can be expressed per
hectare as a parameter of supply of organic matter to the soil.
2. Number of seminal andnodalroots. Particularly in monocotyledons. The
number of nodal roots depends on soil properties.
3. Number of branch roots anddiameter.Absorptive capacity depends on
branching and diameter of the roots.Root diameter can help indicate the
comparability of two root systems.
4. Verticalquantitative distribution in the profile. The amount of root in
different layersis anindication of the nutrient and moisture status of these
layers, and the crop's resistance to drought.
5. Lateralquantitative distribution in the profile. Lateral distribution has a
bearing on such tillage operations as mechanical hoeing and on plant
spacing.
6. Maximum depth and extent. Maximum depth indicates the limit of the
plant's activity, in taking up nutrients and moisture. The significance of
extent (maximum lateral spread) depends on the layer in which it occurs.
7. Courseof development. Observations at different growth stages indicate
rate of growth. Drought or frost are withstood better if plants have esta-
blished agood root system early. Rapid root growth may also counteract
leaching of nutrients.

1.2. Typesoftechnique

1.2.1 Methods infield trials

Monoliths (Chap.2).Thepinboard method gives a fairly complete picture
of the structure and shape of the root system and of the total amount of
root. It supplies data on the distribution in the profile and maximum depth.
Course of growth and branching can be assessed from pinboard specimens
taken at intervals.
Soilcores(Chap. 3).Fragments of root in small soil samples are taken by
augers. This method gives accurate data on the amount of root in each
layer. This is also afairly simpleway of obtaining information on vertical
distribution and maximum depth. Samples at various distances from the
plant show lateral spread and differences between samples taken at inter-
vals show growth rate.
Fieldassessment (Section 3.5) is a simplified process developed from the
auger method.
Excavations(Chap. 4).
Profile walls (Chap.5).This method includes quadrat mapping and root
counting. It is avery suitable method of determining root distribution in
the profile and maximum depth. The total amount of root in any quadrat
can be counted. The method can easily express root branching and growth.
Besides quadrat mapping, roots on the wall can be cleared of soil and then
drawn or photographed (Weaver, 1926).
1.2.2 Methods in container trials (Chap. 6)
Plants can be grown under controlled conditions in containers or combina-
tions of containers. Experimental design and analysis of such trials is often
simpler than of field trials.
The container may be earthenware or asbestos pipes, concrete containers or
glass-panelled boxes for soil cultures and glass cylinders for water cultures.

2. Monoliths: the pinboard method

Rotmistroff (1908) used this method for plants grown in boxes. Maschhaupt
(1915) was the first to use it for a field crop. Later Goedewaagen used it
for plants grown in boxes and in the field.

2.1. Equipment
To construct a pinboard two similar-sized pieces of plywood may be used .
1.5 cm and 1cm thick. In the board 1.5 cmthick small holes are drilled in
vertical and horizontal rows 5cm apart to hold pins made from cut-off
knitting needles or stainless steel wire bent into a Uwith a 5cmbase and
uprights whose length is the sum of the required length of the pins plus the
thickness of the board (de Roo, 1957) (Fig. 1).The piece of plywood 1cm
thick is screwed on as backing.This shape of the pins prevents them from
pushing through the back of the plywood when they are driven into the soil.
If, as formerly, the pins are driven directly into the board it is advised to
use a small tubewhose length is equal to the required pin length. This tube

upper 1.5
board ^_

Fig. 1 Constructionofpinboards
 Fig.2 Pinboardsofvarious
sizes

has a flat base. The tube prevents the pins from going crooked.
The dimensions of the pinboards and length of the pins differ according to
the plants to be sampled (Fig.2).The two most common sizes of boards
are 1m x40 cmwith pins 9.5 cmlong, and 1m x 60 cmwith pins 15cm
long.Thefirst type is for plants with restricted root systems, e.g. pasture
plants and cereals, and the second for crops with extensive root systems,
e.g. potato, beet, rape. Before sampling black polyethylene sheeting is
stretched over the board and pressed with alath down into the pins until it
is against the board. The advantage of polyethylene sheeting is that the
root system can be removed from the board intact after washing off the soil
and draining.

2.2. Obtainingpinboardsamples
When the pinboard is ready the plants are inspected. The crop stand and
development, and any weed growth are noted. Alongside a representative
site, apit is dug about ametre square and the vertical wall against the
plant or plants is smoothed off with a blade sliding up and down two
supporting vertical laths (Fig.3).
When plants are in rows, the pit is dug either parallel to the row'or trans-
versely to it. The specimen taken parallel to the row will contain part of a
*single row as wide as the pinboard and the specimen taken transversely
 Fig.3 Bladeforsmoothing
aprofileface

will contain plants from two or more rows depending on the width of the
board. With a section across two or more rows differences in root distribu-
tion in and between rows, and the lateral extent of roots can be examined.
If the wall is parallel to the row, the distance of the plants from the wall
should be a few centimetres, so that the plants come to lie halfway up the
pins or slightly beyond the middle of the soil slice after the monolith has
been cut out. The soil profile is described and all relevant data are noted,
e.g. type of soil,watertable or height of the site above the ditch level, and
thickness and characteristics of the surface soil and subsoil.
The force required to drive a conical object into the various soil layers with
a penetrometer is measured. At constant bulk density this force increases
asmoisture content decreases, and at constant moisture content it increases
with bulk density (Schuurman, unpublished). Observations are noted on a
standard form. If necessary, soil samples are taken in rings 5 cm diameter
and height to the laboratory to determine bulk density. Roots from the
profile face are teased out with a probe or the face is scraped with a three-
tined hand cultivator to gain an impression of depth and extent of rooting.
Following this the board isheld vertical with the pins against the profile
face so that the top row of pins is at ground level. If the surface is uneven,
the highest point is selected. Thepins are driven into the soil with a jack
(Figv4) or byhammering the back of the board with awooden or rubber
mallet. These operations over, large plants on the pinboard are cut and
stored in plastic bags to avoid damage and loss of water.
After pressing the pins home, a few centimetres of soil underneath the
pinboard is cleared, to a distance of a few centimetres beyond the tipsof
the pins (Fig.4).Thepinboard is then supported by the jack. Soilfrom the
profile face is cut away on either side of the board, also afew centimetres
further than the tips of the pins.Asteel cable,up to 2mm diameter with a
handle at each end, is passed, down one side, along the bottom and up the
other side of the pinboard, and drawn up and down in a sawing movement
so that the monolith is cut awayfrom the soilmass.When it is free, the
pinboard is lowered, still resting on the jack, until it lies on the pit bottom.
The soilis now held on the pinboard by the pins.
Two or four men can hoist smaller specimens out of the pit, if necessary
with ropes.For heavy specimens, use of rope, block and tackle is advisable.
The tackle should be fastened to the back of the pinboard before sawing off
the monolith. Two smooth beams,resting in the pit bottom under the pin-
board are leaned against the top edge of the opposite wall of the pit. The
monolith can then easily be lifted along these beams, for instance, by
towingwith a car.
If horizontal pinboard specimens are needed (Fig. 5),the aerial parts of the

pinboard

jack-

• - f l f c

'%/////////W//////)W/M//.
Fig.4 Useofajackduringpinboardsampling

7
Fig. 5 Horizontalpictureofarootsystemonapinboard

plant are first cut off. The surface is levelled, if necessary.The pins are
pressed into the soil from above.The soil is dug away round the board and
a wooden frame is fitted round the pinboard and specimen. There is a
groove in the frame below the level of the pins.With a jack, a steel plate
with a triangular cutting edgeis forced under the specimen. To prevent the
specimen from being dislodged, the other side should be held inplace.
Once the specimen has been cut loose,it rests on the metal plate and can
be lifted out.Allthe specimens arewrapped in sacking to prevent them
from breaking during transport to the laboratory.
Fig. 5illustrates the extent of a root system of potatoes in the topsoil.

2.3. Removingsoilfromtherootsystems

2.3.1. Sandyorloamysoils
If the monolith consists of sandy or loamy soil,it is directly placed in a
slightly inclined bath of internal dimensions 150x95x35cm (Fig.6).The
lower end of the bottom has 4 plugholes for quick drainage and is placed in
a drain. The lowervertical wall of the bath has a series of 4 plugholes too,
for maintenance of water levels of 3, 10, 17 or 24 cm (Fig.7).The specimen
is put in the bath, then the bathisfilled withwater and the specimen is kept
immersed until the soil seems saturated, usually 12 to 24 h. Clods are then
Fig.6 Viewofwashing baths

Fig.7 Detail ofwashing bath

9
lesslikely to break off and roots to be lost. Washing can then begin.
Damage during washing is avoided bykeeping the water level in the bath
just below the top of the pinboard specimen. This precaution is not essen-
tial. Sandyor loamyspecimens can be washed with ahand-sprinkler or
various types of automatic sprinkler.
Hand-sprinkler.One person can wash the specimen with a hand-sprinkler,
but few technicians keep the pressure constant, particularly if there are
harder and softer layersin the profile.
Rotarysprinkler.The specimen can be washed with a rotary sprinkler with
three arms 20 or 33 cmlong and perforated underneath slightly off-centre
with arow of small holes (Fig.8).The number of arms can be reduced to
one byplugging the other ones, so that water pressure can be regulated.
The slightly oblique position of these holes causes the sprinkler to rotate
with the pressure of the water. The drops scatter evenly over the entire
trajectory.
This method has several advantages. Allmaterial is washed with the same
force. Onetechnician can simultaneously supervise the washing of several
profiles. Such debris as pieces of straw, stalk or leaves should be removed
during washingwith tweezers before they become entangled in the root
mass. If washingis properly supervised, the amount of water need not

Fig.8 Rotarysprinkler

10
 Fig.9 Oscillatingsprinkler

exceed that used with a hand-sprayer.

It is general rule that washing should not be too vigorous since ahigh
pressure may damage roots, and wash away cut roots from plants outside
the specimen.
Oscillating sprinkler.Arectangular area of the specimen may be automati-
callywashed with a modified garden sprinkler placed on a stand. The nozzle
moves to and fro with the pressure of the water (Fig. 9).This area depends
primarily on the distance from the nozzle to the specimen and the length of
the nozzle.Therefore the construction of the stand allows the sprinkler to
be placed at three heights. In each position the spray can be switched to
broad, narrow or to one side.Apinboard specimen size 60x100 cm can
sometimes be washed in a single operation. The apparatus has the advan-
tages of the rotary sprinkler but is superior since it cleans a rectangular
area. It can alsobe used to compare the hardness of layersin two profiles,
since apart of both profiles can be washed simultaneously and hence with
the same force. During washing debris also should be removed by tweezers.

2.3.2. Heavy clays

Pinboard samples of heavy clay are extremely difficult to wash, sometimes
it is even impossible without special preparation of the sample. Attempts
have been made to use amotor pump for washing with aqueous solution of
sodium pyrophosphate (Na4P2Û7).The solution was pumped from a tank,

11
sprayed on the specimen, drained to the tank and recirculated several
times. These experiments were unsuccessful. Fairly good results have some-
times been obtained by drying the entire specimen at 100°C and then
soakingitin an anhydrous sodiumpyrophosphate solution, 270gram per
100litre.The drying allows the solution to penetrate quickly and evenly
into the soil.Drying should begentle to avoid severe cracking, which will
break roots.After about 12hoursin the solution an attempt can be made to
wash the specimen in the usual way.If the sodium pyrophosphate has not
penetrated enough the processis repeated. Recently amethod has been
developed for processing soil cores (Section 3.3.2) byfreezing and action of
sodium pyrophosphate under vacuum. It has given also favourable results
with smallpinboard samples,but the latter must bekept longer than the
auger samplesin the vacuum chamber (Fig. 32).

2.4. Finalsteps
After the specimen has been washed the pinboard and roots are transferred
to alevelbath of water. Here the roots are profiled and cleaned. During
profiling, dislodged and loose roots twisted round needles are restored to
their original position. In the cleaning process debris, e.g. straws, leaves
and roots of adjoining plants, etc., are removed. The specimen may be at
least partially cleaned, byfilling the basin with water and then passing
through it a thin stream of water which runs over the edge.

Fig.10 Woodencasewith
lightbulbsusedforphoto-
graphyofarootsystem

12
2.5. Photography

2.5.1. Root systems

After the specimen has been washed and arranged the root system can be
photographed under water asit lies on the pinboard. Under water the
natural position of the finer roots is also preserved.The black polyethylene
sheeting makes a good background. Roots can be evenly illuminated in a
boxin which are arranged a number of bulbs or fluorescent tubes (Fig.10).
Thebulbs or tubes are placed sufficiently obliquely to avoid reflexion within
the area of the sample. Since the photograph is taken vertically the front of
the camera should be blackened in order to prevent it from being reflected
in the water. Ceiling reflexion may be avoided in the same way. Alternative-
ly four powerful 'Fotomirenta' bulbs can be arranged round the bath to

Fig.11 Submergedroot
systemonapinboard

13
 Fig.12 Submergedroot
systemofFig.11removed
fromthepinboard

give a good distribution of the light and to avoid their reflexion within the
area of the root system.Figure 11shows a root system photographed on a
pinboard under water. This figure illustrates the drawback of pins on a
picture.To prevent this the root system on the polyethylene backing can be
transferred to another bath and then photographed (Fig. 12). See also
Section 2.8.3.
Athird possibilityis to photograph the dry root system after it has been
arranged. Thewater is carefully siphoned out of the bath until the roots
arefree of the water. Theroot system is dried with afan until the outside
is practically air-dry. Theroot system and the polyethylene sheeting are
carefully lifted off the pinboard and the root system is transferred onto
black velvet or black-painted hardboard, for photography. The finer roots
are not so clear as when photographed under water (Fig.13).Moreover,

14
 Fig.13 RootsystemofFig.11
photographedindriedstate

later description of roots is practically excluded. Thebest results are usually

obtained by the second method.
It is clear that pinboard samples of larger plants do not include an entire
root system but only a sort of vertical cross-section of it. Roots towards the
front and the rear of the specimen are cut off.
However, a pinboard specimen gives a good impression of the root system
as awhole.Hudig (1939)moreor less circumvented this limitation by taking
a cube of soil on two pinboards size 30x30cm at right anglesto each other.

2.5.2. Shoots
The aerial parts of those plants that are not cut off are photographed with
the roots. Largeplants are photographed separately, if possible from the
same distance asfor the roots so that photographs are on the same scale.

15
 2.6. Calculationofrootweights
After photographing the root system, the roots can be collected either as a
whole or in layers to estimate the weights. Layers may correspond with
soilhorizons or be chosen arbitrarily. Roots are dried at 75°C before
weighing.

2.7. Descriptionofrootsystems
Aroot system must be considered as a whole before describing individual
rootsin detail.
Ascheme for description is given in Appendix 1.

2.8. Description ofindividualroots

2.8.1. Selectionof roots

For detailed description of roots, e.g. their branching two or more fully
grown roots are selected and worked loosewithout breaking off branch
roots. Thisis a difficult task that requires skill.Experience has shown that
the method succeeds evenwith dense root systems if no force is used. The

cm
20

roots
cm
0

20 20

40 40

60 60

Fig.14 Rootsofoneoatplant

16
 number of roots required depends on the type of investigation and the kind
of root system. Some plants such as potato have roots, differing markedly in
growth habit, one part being confined to the top soil and another growing
in a distinctly vertical direction. Roots have to be taken from both groups
for detailed investigation. In the case of cereals either the nodal or seminal
roots, or both canbe taken.To analyse the effect of soil factors it is useful
to choose fully grown roots.The oldest roots are better avoided as they may
bepartially decayed. This method also affords agood idea of the growth
of the roots of different ages of aparticular plant byworking loose roots
from young to old (Fig.14).
For selecting roots the best procedure is to place the root system in along,
fairly broad and shallow bath with 4to 5cmwater. The bottom of this
bath ispreferably dark for pale roots and light for dark ones. Since the bath
is wide, anyroots not used can be put aside without being cut off at once.
However, the best procedureis to start with cutting awayyoungunbranched
roots since this makes it easier to free fully grown, densely branched roots.
Roots selected for description and photography are cut when they are freed.

2.8.2. Preparing roots for photography

If the root is not needed for weight determination it is transferred to along,
comparatively narrow and shallow bath the bottom of which is lined with
a sheet of fairly firm transparent polyethylene sheeting without folds or
creases asit may later be used for photographing or photocopying. The

Fig.15 Arrangementofrootsbymeansofpins

17
margins of this plastic sheet are weighted to prevent it from drifting when
the vesselis filled with a thin layer of water. If the water is too deep,
fumbling may cause water currents that displace the roots. Fortunately
when roots have only afew side-roots the water adhering to them is suffi-
cient. The branch roots are arranged with pins (Fig. 15) from the baseof
the main root up to the tip.When the whole root has been arranged, the
water is carefully removed with awater-jet vacuum pump. To prevent the
roots from being displaced bythe water they are weighted with glass rods.
The roots are then left to dry until they begin to adhere loosely to the poly-
ethylene sheet. Toprevent roots coming awayfrom the sheeting, as happens
with uneven drying avaporiser is used for spraying the roots with a clear
glue: Saba 810E,dissolved in water in aratio of 1:5.Excess glue can be
removed with awater-jet vacuum pump. The glue takes about 30 minutes
to dry, or less with a fan. The roots then stick to the polyethylene sheet.
Afterwards the dry specimen is covered with a second sheet of thin poly-
ethylene to protect it (Fig.16).

Fig.16 Rootspartlybetween
polyethylenesheets

18
2.8.3. Photography
Before photography of the roots the thin polyethylene cover which could
create reflexions should be removed. Roots can be photographed by either
transmitted light or with ablack background by reflected light (Fig. 17).
The roots should again be covered as quickly as possible with a thin sheet
of polyethylene.

Fig. 17 Woodencasesuitableforphotographywithtransmittedlight

Fig.18 Seminalrootofoats(photographedwithtransmittedlight)

19
 The thin polyethylene sheet is also removed when photocopies are made.
Thelight-sensitive paper comes into direct contact with the roots and a
very sharp photocopy is obtained of the thin roots. Rootswhich vary exten-
sivelyin thickness make poor photocopies and are better photographed.
Photocopies or photographs, which give an accurate picture of the roots,
can be used afterwards for measuring and counting.
The arranging of roots could also be carried out between glass sheets,
instead of polyethylene, but then onlyphotographs can be taken since
these cannot be used in a rotating photocopier.
Roots needed also for weight determination are arranged in a shallow
water layerin aglassbath (Fig. 17).They are photographed in this bath
with either transmitted or reflected light (Fig. 18).In the latter case a
black background of polyethylene is used.

2.8.4. Description
For measuring and counting, the root or a photocopy is laid under a bino-
cular microscope with a calibrated background'(Fig. 19).The binocular
microscope can be moved from side to side and backwards and forwards
independently. Alternatively branch roots are counted by projection on a
squared screen. The description schemes used are given in Appendices 2
and 3.

Fig.19 Binocularmicroscope

20
2.9. Specialapplications and difficulties
If the pinboard is too short for the root system to be sampled, two pin-
board specimens can sometimes be driven close together one below the
other into the profile face. The specimens cannot be cut away until both
pinboards are in position and the soil between them has been cut carefully
with a knife to avoid tearing of the roots.
It is possible to supplement the data with boring specimens taken from the

Fig.20 Rootsystemofoats
andcorresponding profile

21
soil of the hole or its immediate vicinity.
To allow comparison of the root system with the soil profile, both the pro-
file and the root system may be preserved. In these cases the sample taken
is slightly thicker than the length of the pins.The soil outside the pins is
used for making a soil film. This method has been described (Schuurman,
1955) but has since been simplified. After drying the root system is now
transferred from the polyethylene sheet to a blackened hardboard to which
roots and monolith are stuck with a clear glue (Fig.20).
Summarizing it will be seen from the above that the pinboard method has
many possibilities. These specimens can be used to obtain information on
all the points listed in Section 1.1. It is also extremely important that it
gives anidea of the habit of the root system of aparticular plant. Moreover
the method is usually fairly easy to carry out and is not very laborious. One
drawback is that quite abig pit is needed for taking apinboard specimen.
In experimental plots, unless the site adjoins a border, the excavated soil is
difficult to dispose of without damaging the crop.The soil is usually
deposited on alarge sturdy canvas sheet that is spread near the pit. Top
and subsoil and if necessary special horizons are kept apart. Moreover use
of this sheet restricts damage since the soil can be tipped back easier.
Despite this the crop is often damaged over an area of about 4 x 3 sq.m.
Usually this upsets yield estimates from small plots. In such cases it is
generally inadvisable to take more than one sample per plot. Even though
the soil layers are kept separate during excavation the profile will never be
entirely the same as before, upsetting results of trials in later years.
Arestriction is also that the method cannot be used in stony soils.

3. Soil specimens having a slight volume: the auger method

3.1. Equipment
The auger method is used both in field trials and certain container experi-
ments. Two types of augers are used. One, the heavy auger, is specially
designed for sampling heavy soils and hardpans. This is driven into the soil
with a mallet. (Fig.21d).The light auger (Fig.21a) was originally developed
for sampling light soils, but after the cutting edge had been serrated it also
proved suitable for heavy soilswhere it is now evenpreferred to the heavy
one (Fig.22).In stony soils the heavy auger must be used. Both types were
developed by Goedewaagen from augers used byVisser, 1943 (cited by
Goedewaagen, 1948).

22
 Fig.21 Augertypeswithtools

Both augers consist of a cylindrical tube with an inside diameter 7cm.The

height of the tube of the light auger is 15 cm and of the heavy one 25 cm.
Formerly augers 4 cmin diameter were used but the frictional resistance of
the inner wall was so much greater on some soil types than in the 7cm
auger, that soilwas partly forced away. With the 7cm auger this occurs so
rarely that it can be discounted. This has been proved bymeasuring the
bulk density. Nevertheless soil cores drilled with the 7cm auger vary in
length, either caused by compression or expansion of the sample in the
auger tube.Therefore the amount of root present is still that from the
original volume of soil despite compression or expansion during drilling.
Ahollow shaft is fixed above the tube.There are marks on both the tube
and the shaft at 10 cmintervals.The light auger has ahandle fixed to the
top of the shaft to enable it to be driven in the soil and pulled out again.
The heavy auger is provided with a striking head in which can be inserted a
sturdy crosspiece that can be used as a handle to lift the auger out of the
soil.The auger may be lengthened by unscrewing the striking head and

23
 Fig.22 Augerwithserrated
cuttingedge

inserting an extension shaft (Fig.21b,c).

The shaft of the light augerisnormally 70 or 110 cmlong so that they can
be used for drilling to a depth of 80 or 120cmrespectively. The short auger
is used for shallower sampling asin this case the operator's weight is put
to better use.Thelong auger is used for deeper drilling.
The shaft of both auger types contains arod with a disc at the bottom
acting as aplunger. Thelight augeris provided with a handle at the top of
the rod.Therod and stamp areused for forcing the soil specimens outof

24
the tube. In the shaft of the heavy auger a short rod is fixed at the bottom
end to a plunger. The other end of the rod is a rack that engages apinion
in a housing about 40 cm above the tube mounted on the shaft. (Fig.21d).

3.2. Sampling

3.2.1. Thelightauger
Byslowly twisting the auger to and fro in short turns it is pressed vertically
into the soil up to the first mark (Fig.23).During drilling the plunger is
forced upward bythe sample.When the auger has reached the required
depth it is rotated several times to free the sample and then pulled out. The
hole is slightlywidened by rocking the auger. This prevents soil and roots
from being sliced off the wall when inserting the auger for the next sample.
The sample is forced out of the tube by turning the auger upside down and
pressing the handle of the plunger with one foot down. It is collected in a

Fig.23 Drillingasample
withalightauger

25
 cardboard dishwhere it can beinspected. Byarranging the samples in
order of depth a rough description of the profile can be made (Fig.24).

Fig.24 Soilcoresarrangedaccordingto
depth

26
Afterwards each sample is transferred to anumbered paper or plastic bag.
The auger is then replaced in the borehole and drilling is continued up to
the next mark. Thework proceeds until no further roots are encountered in
the sample but, to be on the safe side, one more sample is taken. If the
build-up of the profile is such that a distinct difference exists between two
layers in one 10 cm sample, this may be subdivided into two parts at the
interface between these layers.This subdivision is usually done.The 0-10
cmlayer of grassland is usually subdivided in this way into layers of 0- 5
and 5-10 cm.

3.2.2. Theheavy auger

This auger should be of a much heavier design, since it is hammered into
the soil with awooden or rubber mallet. To prevent damage to the shaft
the auger should be hammered gently. In heavy soils it is often impossible
to push the sample out of the auger by hand. This is therefore done by
rotating the pinion in the housingwith a crank (Fig.25).After that the
same procedure can be followed as for the light auger. In clay soils the
work, with either the light or the heavy auger, may be greatly facilitated by
briefly dipping the auger into apail of water before each drilling operation.

Fig.25 Removingthesamplefromaheavyauger

11
3.3. Calculation of root weights

3.3.1. Handwashing
Drilling samples can be used for calculation of the root weights, after the
specimens have been washed in the laboratory. Sometimes, however, they
have to be prepared for washing. The procedure depends on whether the
samples can be processed immediately, whether the moisture content has to
be estimated andwhether the samples are sandy or clay. Sand samples
processed immediately on arrival and requiring no estimation of moisture
can be washed at once.If the samples, irrespective of the soil of which they
consist, cannot be processed immediately, they are dried at about 100°C
and stored to prevent the roots from rotting. If the moisture content has to
be estimated the samples are packed in the field in heat-proof plastic bags
and areweighed in these bags,before being dried.After drying they are
weighed again. Clay samples cannot be washed until they have been first
dried and then the clay dispersed in a sodium pyrophosphate solution.
Special care should be taken when washing dried samples. If dry fine roots
were washed immediately they would be reduced to powder and lost.To
prevent this, dry samples of sandy soil with the field data on alabel are
soaked in large bottles of water for about eight hours.About 5ml deter-
gent in 300 ml of water assists soaking. During this period the roots also
take up moisture and become so flexible as to enable them to be washed.
Claysamples should be soaked in an aqueous sodium pyrophosphate solu-

•H^BBHHBBBHHE5?1MW Fig.26 Viewofwashingtable

28
tion (270gNa4P2Û7 in 100litre),to disperse the clay.As a result the
sample often disintegrates altogether. (Fig.31a).
Although the roots are fairly shrivelled after the samples have been dried,
they swell again when the samples are again contacted with water. The
roots then substantially regain their normal habit, and even the root hairs
do not seem to be greatly affected. This means that samples first weighed
moist and then dried and again weighed to estimate bulk density, pore
volume and the soil-water-air ratio, can easily be wetted again and washed
for a qualitative and quantitative evaluation of the root fragments in the
sample. This method can therefore be profitably used in studies on the
influence of chemical and physical properties of soil on root development.
The sample iswashed bypouring it out through copper gauze about 0.3 mm
gauge on a specially designed table (Fig.26).These meshes are so fine that
hardly any roots are lost. The gauze usually lets through most of the soil,
except for clods and coarser soil components. During washing free roots
are picked off from the gauze with tweezers and transferred to small bottles
of water ready to hand (Fig. 27).Unfortunately not all technicians maintain

Fig.27 Handwashingofasoilsampleonascreen

29
m)^m^--~n
Wit'î'

»fi«»!;,,
"""SB?

;?r
"ç^l^#.

Fig.28 Rootsstoredinabottleandinapolyethylenebag

a constant water jet, and results may vary between workers.

When no more soilpasses through the screen and onlyveryfine roots are
left, everything on the screen -rootlets, humus components, plant debris
and soil particles -iswashed into a glass cylinder in which the soil particles
settle out and roots and other organic material continue to float awhile.
Bydecanting once or more and topping up the roots can be separated from
the soil, but anyplant debris and humus components remain among the
roots, as they cannot be separated from them by decanting. Finally the
mixture is poured onto a fine nylon or muslin gauze measuring 10x10cm
whence the mass remaining on the gauze is added to the other roots that
were already separated.
If the samples cannot immediately be dealt with, formalin or ethanol 60%
or 1.5 gram finely divided thymol per litre is added. Since glass bottles are
breakable and take up space, roots have recently been packed in plastic
bags with some water, heat-sealed and frozen (-20°C) or with some pre-
servative solution (thymol) and then heat-sealed (Fig.28).Afterwards the
organicimpurities are also removed. This is done at a specially designed
table (Fig.29) bypouring the sample into a shallow enamel bath measuring
26x20x4cm, after which the impurities are removed with tweezers (Fig.
30). If there are manyimpurities removal of the roots from the mass is
better.

30
 Fig.29 Viewoftable for
separating roots and debris

Fig.30 Separation ofroots and debris

31
It is difficult to distinguish living and dead roots, i.e. roots which were alive
or dead when sampled. Four features are noted for drawing this distinction:
the elasticity of the root, its colour, and the presence of cortex and lateral
roots. Adead root is far less elastic, is often grayer, the cortex is often
ravelled or it has no cortex and lateral roots have often broken off, leaving
stumps with frayed ends.The combined assessment of these four characte-
ristics determines whether a root should be regarded as alive or dead.If
dead it is removed.Another test, although unsuitable for routine is to
contact the roots with atetrazolium chloride solution. Liveroots stain red,
whereas the dead ones remain colourless (Goedewaagen, 1954; Butijn, 1955
and 1961).After removing allimpurities the roots are again tipped out on
to the fine gauze,moisture is gently pressed out and then the roots are
transferred with tweezers to a smallpaper bag on which all data are noted.
The original label is also placed in the bag. Finally the roots are dried in the
bags at 75°C. It was found, that the results hardly differed from those
obtained at 105°C. Afurther advantage of this'lower temperature is that it
prevents the roots from being pulverised. After drying for about 48hours,
the bags of roots are placed in a desiccator to cool.Theroots are then
weighed.
The intention of washing is to remove all soilparticles.They can, however,
never all be removed as some particles adhere firmly to the roots, especially
if the roots have an abundance of root hairs. The presence of soil particles
biasses the root weights obtained. This inaccuracy was checked periodically
by occasionally ashing washed and dried root samples. The percentage
inaccuracy usually increases the deeper the layers from which the roots
were taken. Themaximum inaccuracy found was 12%.Fortunately only a
small proportion of the roots is found below a depth of about 20 cm. It can
be stated as ageneral conclusion that with carefully washing the impurities
need not exceed 4%, for the entire depth of sampling. In some cases even
lower values have been found. If avibrator is used after washing this per-
centage can evenbe further reduced.As arulewe didnot estimate organic
matter contents of the roots, although it can be done with the auger
method.

3.3.2. Routine method

In the previous pages a simple method for processing soil cores obtained by
augers has been described. Even though the method maybe considered as a
good one it is clear that the operation contains personal elements, for
instance when not everybody works with a uniform water yet. Moreover
the method is very time consuming. Finally drying of soil samples, however,

32
necessary to preserve them, may be destructive to the roots.
Wehave developed a routine automatic and standard method that complies
with reasonable requirements of being objective, not laborious, gentle so
that the soilparticles are loosened from the roots without damage to the
latter, and simple provided the necessary equipment is present.
Firstly, soil samples, having normally avolume of 385 cm3 are thoroughly
frozen in the laboratory at a temperature of -20°C. While they are thawing
they are submerged in about 800ml solution of sodium pyrophosphate. The
favourable concentration of sodium pyrophosphate after the freezing is
2 to 4grams per litre (Fig. 31a). In contrast with this the action of sodium
pyrophosphate without freezing and vacuum is small (Fig. 31b).No distinc-
tion is made between clayey and sandy soils.The samples are then placed
in avacuum chamber. When a small number of samples are to be processed
an undamaged andhigh quality desiccator can be used. For many samples
proper equipment is required (Fig. 32).The first step is then to evacuate
the chamber to apressure of about 45 cm Hg.Following this, air is admit-
ted above the solution surface to restore normal pressure. Then again air is
evacuated, now until the pressure is about 25 cm Hg.Air bubbles show that
air is escaping from the sample. Again normal air pressure is re-established.
Finally air is pumped out until a pressure of about 6 cmHg is attained and
this is maintained for about 15minutes. Following this, air pressure is
restored again to normal. Bythis alternating from a vacuum to normal air

J.J 3.5 4
mmm
4.5

1 1.5 2 2.5 3 3.5

•III
4 4.5 5 0
Fig.31 Effect ofvarioussodiumpyrophosphateconcentrationsuponthesoil.
A. undervacuumafterfreezing; B. withoutvacuumandfreezing

33
 air ouüet for lower
chamber
inlet
pyrophosphate

Fig.32 Drumfortreatmentofsampleswithsodiumpyrophosphateunder vacuum

pressure the sodiumpyrophosphate solution penetrates deeply in the soil

sample and causes anearly complete disintegration of the soil.This means
that the roots are practically completely separated from soil particles
although they are still mixed with these.The sample is next transferred to
the top compartment of aplastic gutter, with a total length of about
V2 metre, divided into 3compartments, separated by removable partitions
(Fig.33).The roots are washed down towards the final compartment by a
stream of water to separate them from the soil particles. During passage
they are picked upwith tweezers by a supervisor and transferred to a gauze.
Roots that pass the gutter unseen are caught in a gauze filter at the endof
the gutter. These roots are mostly mixed with plant debris.Bydipping the
gauze in about 3cm of water usually the roots and debris can easily be
separated. Thewashing operation takes 5-15 minutes. One technician can
supervise at least 8gutters simultaneously. Treatment with sodium pyro-
phosphate and washing of about 25 samples takes about 60minutes. It is
calculated that one person can process between 100 and 150 samples per
day.
If soil particles remain sticking to the roots after processing, the roots can
be put in small bottles of water that are placed one byone on a vibrating
apparatus for up to 3minutes.This almost completely separates roots from

34
Fig.33 Gutterfor separationofrootsandsoilparticles

sand particles.
Themajor part of the roots is recovered practically undamaged (Mesker and
Schuurman, in press).

3.4. Results
The auger method supplies only data on fragments of the root system and
givestherefore little idea of the entire root system as awhole.Nor is it
possible to study the branching because many lateral roots are wholly or
partly cut off bythe auger. Onthe other hand, samples can be taken for
weight determination without damage to the experimental field. Moreover,
periodic auger samplings can indicate the development of the root system
at particular sites.Due to the variation in root weight in the various soil
layers an accurate idea of the roots weights can only be obtained if a large
number of samples is taken from which the average values per layer can be
calculated. The statistical reliability of these figures can then be deter-
mined. Anexample is given in table 1.
In our studies the minimum number of borings per plot of mixed cultures
as grassland was set at 25; at least 24 borings are taken of such mono-
cultures as arable crops. Where such crops are cultivated in rows, e.g.

35
Table 1. Reliability of root weights from auger samples of a pasture

Depth(cm) Weightof Weightof ó m mo

25root rootper
samples sample

0-5 3673 147 65 13 8.

5-10 1112 44 14 3 6.8
10-15 917 37 14 3 8.1
15-20 483 19 9 2 10.5
20-30 544 22 9 2 9.1
30-40 327 13 7 1 7.7
40-50 201 8 5 1 12.5
50-60 158 6 4 1 16.6
60-70 66 3 2 0.4 13.3
70-80 73 3 3 0.6 20.0
80-90 3 sp 0.4 0.09
Total 7557 302

cereals, 12borings are taken in the rows and 12between. For crops not
sown in rows other sites maybe selected. The holes are drilled over the
field in a selected pattern. Weedy patches are avoided. Since in most crops
the weight of roots in the topsoil is 70% to 90% of the total, an accurate
estimation of the weights of the surface roots is more important than of the
subsoil roots.For this purpose, it is possible to adopt a system of taking a
smaller number of complete borings with an additional number of samples
from the topsoil. Although accuracy in the root-deficient subsoil is reduced
as aresult, there is a slightly greater accuracy in the root-rich topsoil.
Since each boring is divided in layers of 10 cmthick or less, some 250
samples are obtained from 25 complete borings per plot at a drilling depth
of 1metre. These samples have to be processed in the laboratory. This
indicates that this method isvery laborious, and this is its great drawback.
Given the diameter of the auger the root production per hectare can be
calculated from the root weights per boring.This value is aparameter of
the supply of organic matter to the soil.

3.5. Field assessment of root density in cores

Preparation of auger samples for weighing is a time-consuming operation.
For thisreason a quicker method has been developed in which an assess-

36
 0.032

0.32

3.2

32

Fig.34 Somestandardfigures for useinfield assessment of roojt densities

37
ment is madein the field ofthe amount ofrootsinsoil samples.Although
slightly less accurate, this method is adequate forfield work and requires
little or no laboratory work. This method has been fully described in an
earlier publication (Schuurman and Knot, 1957).Here we will only givea
brief description ofits principle. Hitherto the method has been elaborated
onlyforcrops like grasses and cereals whose roots have no secondary
thickening.Itis based on acomparison ofroot quantities with standard
data.
The samples for the estimate aretaken with the auger.After the sample has
been pushed out ofthe auger itis broken horizontallyin the middle.The

root w e i g h t
in kg/ha

bUUU
« .
3000
2000 - • *

1000 - »• ••
- *;.V
500

300

200 * Jr• • •

100 r .• •3&•
' . V• -a«.•'.
Ma« «• »A
• • • •A
• •••j* • M
50 • •I I . — •

30 - •• } •• •

20
• M • •

10
• '
5

3 -
2 -

1
i •• I 1 1 1 1 ,1 1 1 , 1
0.01 0.02 0.05 0.1 0.2 0.3 0.5 2 3 10 20 30 50
root c o v e r a g e

Fig.35 Correlation betweenestimated valuesand ovendryweights of roots

38
roots growing across this plane of fracture do not usually break in this
plane but near to it.They can then be seen and they can be compared with
specially designed standard figures. These consist of a series of circles of the
same diameter as the auger with an increasing number of light dots on a
dark background (Fig. 34).Each circle has a known dotted area. The total
coverage of the roots is calculated by adding together assessments for both
planes of fracture. The accuracy can be enhanced bybreaking the core at
several points and assessment in allplanes.This is especially important for
samples in which there is a sharp decrease of the amount of roots in a
downward direction. It is then possible to calculate the mean coverageof
these planes of fracture. In order to eliminate the 'dissimilar height' factor
of samples, the coverage figure is multiplied bythe height of the samples
expressed in centimetres. It has been found that root coverage is correlated
with root weight of the same sample (Fig. 35) (Schuurman and Knot, 1957).

3.6. Specialapplicationsand difficulties

Samples taken with the auger in wet soils maynot come up when the auger
is lifted out. This can be overcome byproviding a light auger with a leather
or rubber piston above the plunger (Fig.36). Since the piston fits tightly it
has to be gradually raised during drilling.When the auger has reached the
required depth the piston is raised abit further to reduce the air pressure
above the sample in the auger; as a result the sample remains in the auger
while this is lifted out.
Good results have also been obtained with an auger round which is fitted
another drilling tube about 2mm wider than the usual one (Fig.37).When
the auger is lifted air can pass down the outside of the inner tube, so that
no vacuum is formed below the sample.
When samples are taken invery soft soil the borehole does not remain
entirely open.Asimilar difficulty occurs when many borings have to be
made close together and the remaining soilhas to be supported. In such
cases useis made of thin-walled iron pipes with a detachable top-piece
secured to the pipe bymeans of a bayonet closure.The pipe and the top
piece are over 1metre long and have a diameter enabling the drilling tube
to fit in itwith someplay.The auger isplaced in the pipe so that the
handle is inserted in the corresponding cavities of the re-inforced upper
edge of the detachable top-piece (Fig.38).The saw-teeth of the auger then
project some 2mm below the bottom of the pipe. The auger and pipe are
both pressed into the soil to the first mark on the pipe which is ma'de 9.8
cm from the bottom. The auger is then withdrawn, the pipe remains in
position and descends 10 cm more with each subsequent boring until the

39
 nil I—thickness of
piston auger wall 1mm

Fig.36 Auger provided with a

rubber piston

40
Fig.37 Augerwithextratube

full depth has been reached. The top-piece is then taken off and used for
the next boring.Thepipe itself remains in the borehole either until all
borings have been made, or indefinitely if more borings are to be taken
periodically from a profile of a cylinder experiment. Moreover, the pipes
seal off surrounding soil, thus preventing changes in aeration and moisture
content in this soil (Fig.39).This method can also be employed in humid
soils.
Difficulties may also be encountered in the sampling of peat soils. In the
first place the peat may be too loose or so stratified that the auger does not
penetrate. Nor isit possible without preparatory measures to wash out all
the peat from a sample without roots being lost. However, there are indica-
tions that apre-treatment with a 5% hydrogen peroxide solution or a
solution of sodium pyrophosphate has a good effect. Peat is also difficult to
wash off from pinboard samples, although the drawbacks in this case are
less considerable as an idea can be formed of the structure of the root
system from apinboard which is still partly covered with peat. Moreover,
with careful handling the peat can be removed with tweezers.

41
 g roove fo k j ^
a uugq e r h a n d l e ^ ^
attachment

for p i n ^ n - ^ X J-auger
handle
il
—attachment
p i n- ! t•
" ^pin

cyli nder w i t h - —cylinder

gr a d u a t ! ons

fix
m
saw toothed edge
of t h e a u g e r t u b e
Fig.38 Augerwithaccessorycylinder

When taking auger samples a further difficulty may arise which is equally
applicable to the pinboard method. Auger and pinboard methods are im-
practicable in very stony soils.In these cases the method developed by
Weaver (1926) inwhich the root development is studied in a profile wall is
used.
Summarizing it may be stated that both the pinboard and auger method
have their own particular advantages and disadvantages and that the data
not supplied by one method can be supplemented by the other. Conse-
quently it is advisable to employ both methods at the same time.

42
Fig.39 Cylindersinthesoil

4. Excavations

Goedewaagen already used rectangular iron frames to determine root

masses in the topsoil as aparameter of supply of organic matter to the soil.
These frames had a length of 24 cm, awidth of 20 and a height of 30cm
or 30x27x40cm.The bottom edgewas sharpened. He pushed or ham-
mered the frames vertically into the soil, excavated them and washed them
on a screen with a spray nozzle.After drying he weighed the roots.
Recently Dilzused similar iron frames that are pushed into a profile wall.
Samples are taken of the topsoil with aheight of 25 to 30 cm and horizon-
tal dimensions of 20x10 cm.The sample is first saturated with water and
then weighed. Bysprinkling the sample with a standard apparatus during
a fixed time part of the soilis washed away.The remaining soil is again
conditioned to the same moisture content and then weighed. This weight is
seen as aparameter of the soil-binding capacity of root systems.This is
particularly important for crops in relation with the sensitivity of the soil
to erosion. Sincewashing of clay soils is difficult these samples are frozen
on arrivalin the laboratory and then after thawing treated like sandy
samples.

43
 5. Profile walls

5.1. Mapping,followed bycounting

Oskamp and Batjer (1932) developed this method for trees.Afterwards we
slightly modified and supplemented it. In this method either tangential or
radial trenches are dug at a certain distance from a tree and the tips of the
cut roots are mapped in one of the walls of the trench. This is done by
covering the face with a system of squares of, say, 20x20cm (Fig.40).
Nails are therefore driven into the profile wall at 20 cmintervals round the
sampling area. Pieces of string weighted at one end are suspended from the
upper horizontal row of nails, so that they hang vertically along the wall.
Other pieces of string weighted at both ends are suspended over the corres-
ponding nails of the two vertical rows thus forming horizontal lines.The
wallis then carefully scraped with along needle (sack needle with a handle)
to reveal the tips of the cut roots. The point of this needle should not be -
sharp, to avoid cutting the roots.These are mapped, different categories
being distinguished according to thickness.Adifferent symbol is used for
each category (Fig.41).The profile may be sketched in on the map at the
sametime. It maybe asked how the trench should be located with respect

Fig.40 Squaresonaprofilewall

44
 breadth in c m GREEN MANURE
0 40 80

/"... • • •

--•V*
^
• * . '
* *• • • • : *

-.*•• o •' -' •

•*•
o. o,', o'

,'• 'o'i'-
'"'•:« 6.: O0?.W°"
'1*0
'.. • )
• yo'.. b
.0*1
g-ft? .'o ? °".» "..

%'...,.o b ' Ó' °o ó

4
Q •••• °o'
0-' ^•H"
.'•-.,
#" 0
p

• •
o
i
» ; %
° " ' f. '1'.....
•
• ] '1 !
diameter of roots : : Ï2, ° 1/2 - 1 , 1 -5, o 5-10, • 10 mm
Fig.41 Examplesofrootmapsof aprofilewall

Table 2. Comparison of the numbers of roots of apple trees on a grass plot in

tangential trenches at equal distances from the tree

Distance from the tree 200 cm Distance from the tree 150cm
Width of the profile 120 cm Width of the profile 80 cm
(= 1/12 circumference) (= 1/12 circumference)

Numbers of roots Percentages Numbers of Percentages

per trench per trench roots per trench
per trench

Layer 1 2 3 4 Av. 3 4 Av. 1 2 Av. 1 2 Av.

cm Diameter of the roots <y2 ^m

0- 10 0 13 4 0 4 0 10 2 0 3 8 14 11 6 6 6
10- 40 22 29 27 42 30 17 22 15 34 22 33 75 54 25 33 29
40- 70 52 27 75 47 50 39 20 40 38 35 34 82 58 25 36 32
70-100 43 39 52 16 38 32 30 28 13 26 37 51 44 28 23 24
100-150 16 24 25 19 21 10 18 13 15 14 22 5 14 16 2 9
Total 133 132 183 124143 134 227 181

Diameter ofthe roots y 2 -l mm

0-10 1 3 3 0 4 13 10 0 3 12
10- 40 9 12 3 13 9 37 54 10 30 30 8 10 9 24 21 22
40- 70 8 4 15 18 11 33 18 48 42 37 10 15 12 29 31 30
70-100 3 2 3 7 4 12 9 10 16 13 11 18 14 31 37 35
100-150 3 1 7 5 4 12 5 22 12 13 1 3 2 3 6 5
Total 24 22 31 43 30 34 48 40

45
number number
_
s t r a w mulcHI c l e a n s o i l p r e e n manure]] g r a s s '
4 d i a m e t e r > 10 mm
•—tree no.2l
2 °—tree no.3 2

0 ./„-' 0

6 diameter 5 - 1 0 mm 6

4 4

2 ky 2

0 0

25 r» 25
diameter 1 - 5 mm
20 20

15 16
10 10
5 5
/ ^ > >
0 0

25 diameter V2- 1 mm 1 2 5

20 20

15 15

10 10

5 5

0 0

150 j- 150

125 -
diameter <V2 125

100 100

75 75

50 (-' 50

25 25

0 0
10 30 50 70 10 30 50 70 10 30 50 70 10 30 50 70
d e p t h in c m
Fig.42 Comparisonoftherootnumbersoftwotreessubjectedtothesame
treatment

to the tree. Somevariations has been found in tangential trenches, but it is

more or less immaterial on which side of the tree the trench is dug,pro-
vided conditions round the tree e.g. the structure of the profile and the
hydrological regime are uniform (tables 2 and 3).Atangential wall can be
divided into anumber of parts of equal sizewhich may be taken as replica-
tes.Actually this division has already been made by the system of squares

46
during the sampling process. This is important as it means that the root
zones can be compared at different times by moving to other places on the
same circle in successive samplings. The radial trench indicates root deve-
lopment at different distances from a tree. Since in such a map each root
has its own distance from the tree there can be no question of replicates
and this means that the data provided by a radial trench are less reliable.
The reliability can, however, be enhanced by sampling a greater number of
radial walls per tree, or both walls of a radial trench.
Tangential trenches also afford an impression of the root development at
different distances from a tree when they are dug at varying distances from
the trunk (table 4).
The comparability of trees treated the same way has been shown by a trial
with four groups of two apple trees in tangential trenches. Data on root
density in various soil layers, given in figure 42 show a reasonably good
agreement.

Table 3. Comparison of the numbers of roots of apple trees on a straw mulch

plot in tangential trenches at equal distances from the tree

Distance from the tree 200 cm Distance from the tree 150cm
Width of the profile 120 cm Width of the profile 80 cm
(= 1/12 circumference) (= 1/12 circumference)

Numbers of roots Percentages Numbers of Percentaces

per trench per trench roots per trench
per trench

Layer 1 2 3 4 Av. 1 2 3 4 Av. 1 2 Av. 1 2 Av.

cm Diameter ofthe roots < y 2 mm

0- 10 130 161 97 98 122 36 45 30 32 36 104 78 91 36 28 32
• 10- 40 135 80 118 120 113 37 22 37 39 34 87 98 92 30 35 33
40- 70 21 31 54 40 37 6 9 17 13 11 29 41 35 10 15 12
70-100 27 28 24 25 26 8 8 8 8 8 31 29 30 11 10 11
100-150 49 56 25 23 38 13 16 8 8 11 37 31 34 13 11 12
Total 362 356 318 306 336 288 277 282

Diameter ofthe roots y 2 -l mm /

0- 10 10 29 16 16 18 16 32 26 20 24 17 8 12 22 16 19
10- 40 36 28 32 35 33 58 31 52 43 44 36 24 30 46 47 47
40- 70 7 19 10 15 13 11 21 16 19 17 13 6 10 17 12 15
70-100 8 10 4 9 8 13 11 6 11 11 10 7 8 13 14 13
100-150 1 4 0 6 3 2 4 0 7 4 2 6 4 2 12 6
Total 62 90 62 81 75 78 51 64

47
Table4. Comparison of the numbers of roots at three different distances
from the tree

Width of the trench 1/12 circumference

Distancefrom the tree

Diameter of the 2metres 1.50 metres 1.00metre

roots inmm

Straw mulch plot

<% 336 282 272
%-l 75 64 54
1-5 32 36 28
5-10 7 6 6
>10 2 5 4
Grasscoveredplot
<% 143 181 150
%-l 30 40 36
1-5 14 22 16
5-10 4 2 3
>10 1 1 0

From the root map the number of root of each size category is counted per
square.From these data the total number of roots in each layer can be
calculated. The average number of roots per unit of length of alayer and
the statistical reliability may be calculated.
Themapping method can provide important information on root develop-
ment of woody plants. It has, however, a number of limitations. In the first
place it is very time-consuming so that it is impractical to sample more
than one tree in aplot. Secondly, as allwork has to be done outdoors, good
weather is essential unless a tent and artificial light are used. Finally, it
should be pointed out that the sampling depth is governed by the water-
table.This method can also be employed for agricultural and horticultural
crops. Trenches are then usually dug parallel to a row or across one or
morerows.
Avariant of this method used in sandy soil, is that the soil is flushed out
with water instead of being worked loose with aneedle.Aknapsack sprayer
of vaporizer can be used for this purpose. After awall is sprayed with the
finest atomizer the cut roots show up well and can be mapped (Fig.43).
Only a small amount of water is required. This variant is also used by other
scientists.

48
Fig.43 Sprayingaprofileface

5.2. Weaver'smethod
Themethod as introduced byWeaver (1926), referred to on page 42is also
used for the investigation of profile walls, especially in stony soils where
pinboards and augers are impractical.

6. Experiments in containers

In these experiments plants are grown under specific, pre-determined con-

ditions. In some experiments several of the methods mentioned above may
be used, but the procedure in many container experiments/is often so
adapted to root studies that the processing has its own particular character.
The following will be discussed in succession: experiments with cylinders,
cases,boxes andpots filled with soil, and cylinders filled with nutrient
solutions. Ageneral and favourable feature of container experiments is that
complete root systems are available for investigations.

49
6.1. Cylinders

6.1.1. Material
Cylinders of different material and dimensions are used. Concrete cylinders
werefirst used by Goedewaagen (Frankena and Goedewaagen, 1942).The
original, laborious method has since been improved and simplified and new
facilities created. Until recently only large cylinders, made of concrete with
a 30 cminternal diameter and aheight of 100or 125 cm, and small cylin-
ders, made of asbestos with a 15cminternal diameter and a height of
75 cm were used.
The large cylinders are arranged in 2groups in large, concrete vessels, that
are divded in two sections by a concrete wall (Fig.44).Each section can
hold up to nine large cylinders.Asconcrete cylinders have aweight of about
100kg each, they are difficult to handle. Therefore recently large asbestos
tubes have been introduced with the same external dimensions. Besides
being lighter these tubes have the advantage df a larger internal diameterof
36 cminstead of 30 cm and they can easily be cut longitudinally thus
facilitating washing.
For both kinds of tubes care is taken that the diameter is correct and the
tube is circular, since this is essential for uniform filling with soil.The
upper edge of the cylinder is about level with the surrounding surface.A
given soil watertable can be maintained in the concrete containers.The
watertable required does not necessarily have to be constant - avarying

Fig.44 Lay-outofanexperimentwithtubesinaconcretecontainer

50
Fig.45 ViewofanexperimentwithPVCcylinders

level may also be used.

The asbestos cylinders 15 cmin diameter, have recently been replaced by
PVCcylinderswith a 20 cm external diameter and aheight of 90 cm.This
material has the advantage of being chemically inert. The cylinders are
usually placed on trolleys so that the experiment can be either out of doors
or under glass, depending on the weather and the purpose of the experi-
ment. Recently they are placed in a movable greenhouse (Fig.45).

6.2. Largecylinders

6.2.1. Filling
The soil with which the cylinders are to be filled is first screened. The
screened soilismixedwith fertilizers as uniformly aspossible and stored in
plastic bags to prevent loss of moisture. The moisture content is determined
so as to enable a calculation to be made of the amount of soil to be intro-
duced into the cylinder in order to obtain a predetermined bulk density.
This amount of soil is weighed out and divided into portions which are
again stored in plastic bags.Each portion is exactly enough to fill the cylin-
der with a 5cmlayer. Before filling the tube is closed at the bottom with
porous nylon cloth (Fig.46) and secured upon a concrete base of equal
diameter provided with two wireloops by means of a bar and screw (see

51
 Fig.46 Part of cylinder
closedwith nyloncloth,and
basewith loops

Fig.47 Apparatus for

compressing the soilalong the
wallof a cylinder

52
 ÄJ»."«M

Fig.48 Loweringfilledcylindersintocementcontainer

also fig. 56).During each filling operation half a bag is emptied into the
cylinder and then tamped down.Aregular check is made to ensure that the
proper level is reached after tamping. If this is the case the soil along the
edge of the tube is pressed down with anarrow curved iron bar (Fig.47),
so as to prevent growth conditions for the roots being more favourable in
this part than in the rest of the profile. The topmost layer of from 2 to 5
mm is then carefully scraped loose over the entire surface to prevent the
formation of layers (Goedewaagen, 1932 p. 182).The second half of the soil
is then poured into the cylinder and treated in the same way. The whole
procedure is repeated with each portion of soil until the cylinder is full. In
this manner it is possible to build up a great diversity of profiles with
variations in soil type and density. The filled tubes, placed upon the con-
crete bottom are transported to and lowered into the container by meansof
hooks and rope, attached to the wire loops (Fig.48).After placing, the
hooks are freed from the loops. Since the soil is capable of absorbing a
great deal of moisture in the initial stages, the containers are regularly
replenished with water until an equilibrium has been reached. The crop can
then be sown.
Recently promising results have been obtained in an experimental tamping
with a slatted iron ringwith a handle.Tamping is done by one operator
while the tube turns round and the soilis tipped in regularly by a second
person.

53
 Fig.49 Regulationofsoil
watertablewithsyphons

6.2.2. Soilwatertable
The watertable in each section of the containers used in the large cylinder
experiments can be set to various heights with siphons (Fig.49) in a central
vessel that are connected with these sections.
Thewater levelin the containers is checked several times aweek and hence
the corresponding soilwatertable in the profiles also.During and after rain-
fall excesswater is removed bythe siphon offering the possibility of mea-
suring these amounts. If the soilwatertable has dropped it ismade up.The
amounts added and of precipitation are a measure of the absorption of
water by the plants. It is advisable to cover the top of the profile with a
layer of fine gravel of about 2 cm thick to prevent evaporation.

6.2.3.Sampling
There arevarious ways of studying the root development of a crop.It has
already been shown inFigure 39how aprofile in a large cylinder can be
drilled. Experience has shown that at least 6borings can be obtained from
cylinders of 30 cm diameter.

54
 Fig.50 Bladesfor cutting
awaysoilhorizontallyin
cylinderexperiments

Asecond method is to remove all soil from the cylinder layer by layer. This
is done byfirst removing some samples with an auger up to the required
depth and then cutting loose the rest of the soil to the same depth by
means of specially designed blades (Fig.50) and taking it out of the tube.
The drilling cores may be kept separate, e.g. for moisture determination,
and added to the remainder afterwards. The resultant samples may be
washed as discussed in Section 3.3.
Athird method is to lift the tube out of the container together with the
profile with a rope and hooks attached to the wire loops.
To prevent the soil from falling out of the tube this is lifted out whilst it
stands on the bottom. It is transferred to the laboratory, where it can be
laid on top of a steel basin (Fig. 51) or on a steel stand. Ofi this stand the
profile can be turned round horizontally and set at various slopes and thus
be adjusted to the available light and most convenient height (Fig.52).The
crop may or maynot be cut before transport. On the basin or stand the top
of the profile should preferably be lower than the bottom, so that some
water always remains in the tube during washing.This prevents the soil and

55
Fig.51 Cylinder aboveawashing bath

56
 Fig.52 Adjustable standto
holdPVCtubeduringwashing

waterinlet

Fig.53 Washingapparatus

the roots from breaking. However, the profile may slide out of the tubeif
the tube has a slope of more than 30°.Thenylon cloth is^removed and the
soil is washed out of the tube. This can be done with a sprinkler about 80
cmlong that allows either sprinkling frontally or sideways, regulated by a
two-way switch (Fig. 53).Usually it is best to start washing from the bot-
tom of the tube. The presence of many roots in the top layer makes it
difficult to wash the soilfrom the top except at the last stage, after the top
layer has been pierced. Moreover when the profile is being washed from the

57
Fig. 54 Removingdebrisfrom arootsystemafter washing

bottom one can immediately note the depth to which the roots have pene-
trated the soil.Aswashing proceeds the slope can be reduced. In the last
stage of washing the tube is so placed that the top of the profile is highest.
Turning the tube during washing must be avoided, since this may damage
the roots and twist them. When the entire root system has beenwashed free
of soilit is carafully slid out of the tube into a shallow bath with the useof
plenty of water. The size of the bath must be adjusted to that of the root
system. Any extraneous matter is removed in this dish (Fig.54).The root
system can then be photographed. Finally the root system and the indivi-
dual roots can be described in the manner indicated in Section 2.7 and2.8.
One disadvantage of the third method is that no data is obtained on the
moisture content of the soil.Thismay be overcome by drilling samples from
the profile with anarrow drill with a diameter of 14mmbefore the soil is
washed away (Fig. 55).Owing to the fineness of the drill it is not possible
to make a hole deeper than about 60 cm.This drilling also results in slight
damage to the root system.
Afourth method of studying root development is employed by cutting the
tubes longitudinally in two slightly unequal parts. The smallest part is
removed, the larger piece of tube is laid with the.profile in a bath. The soil
can then be washed with an oscillating sprinkler (Fig. 9).This method saves
much work and provides the least damaged root systems. Water content of
the soil should be determined as described in the third method before the
tubeis cut.

58
 Fig. 55 Drillsusedtoobtainsmallsoil
samplesformoisturedetermination

6.3. Small cylinders

6.3.1. Filling
The small cylinders are usually above ground. They are filled with soil in
exactly the same manner as the larger cylinders. (Fig. 56).It is also advis-
able to give the bottom of these profiles apermanent support by securing
moisture-permeable nylon cloth to the bottom of the cylinders. In general
the same kind of experiments can be carried out with these cylinders as
with the large ones. Since they are lighter they are easier to handle, but the
cylinders with a diameter of 15 or 20 cm do not take as many plants.

59
 t e n s i o n bracket

&
rod

wooden
bottom

Fig.56 Woodenboardwithtensionbracket

nylonclcth

Big.57 PVC-tubewithaccessories

6.3.2. Soil watertable

The cylinders are placed in larger ones in which the watertable can be
maintained at various levels (Fig.57).In experiments where only deep soil

60
 c y l i nder wal

rubber rings^

r u b b e r col

polyethylene-
collar
Mi t s c h e r l i c h
dish

Fig. 58 Tubeinadishclosedbyrubberandpolyethylenecollarstoprevent
evaporation

watertables are wanted shallow dishes may be used. To prevent evaporation

from these vessels th?y are covered with a rubber collar. This collar can
also prevent rain from seeping in the vessel when the experiment is in the
open, but as an extra precaution a second collar of flexible polyethylene
material is then placed above the rubber one.This polyethylene collar is
fastened between two strong rubber rings (Fig. 58).Evaporation from the
profile is prevented by covering it with alayer of fine gravel or plastic
granules asis done for large cylinders.Thewater used for replenishing the
vessels is measured, so that the moisture uptake of the plant in each cylin-
der can be accurately determined.

6.3.3.Sampling
The roots are examined bywashing away the soil, as discussed in connec-
tion with the large cylinders. Moisture samples are taken with the small
/
drill (Fig.55).

6.4. Special applications

Hitherto wehave only discussed cylindrical tubes, but it is obvious that
both the shape and material may be adapted to circumstances.
Cylinder experiments canbe carried out with both natural and artificial

61
profiles. In their studies Frankena and Goedewaagen (1942) forced a steel
tube with a cutting edge into the soil.This tube had a diameter of 30 cm
equal to the internal diameter of the concrete cylinders.After being exca-
vated the profile was transferred to a concrete cylinder.
Later experiments showed that it waspossible, and probably even better, to
excavate a soil column with a 30 cm diameter on which the cylinder stands.
Asthe soilis dug more deeply and cut away, the cylinder gradually falls
round the soil column byits ownweight.Ametal collar with a 30 cmdia-
meter and provided with a cutting edge may be used. In this case the cylin-
der rests on the metal collar which it forces downward. This method can
only be used in soils with a firm profile structure, i.e. the soil should
neither be too wet nor too dry. Should it be too dry, one can attempt to
make things easier bywetting the soil.Till now better results have been
obtained with glazed earthenware cylinders than with the fairly rough can-
crete cylinders.
Undisturbed soil samples have likewise been obtained in cylindrical tins
20 cmhigh and with a diameter of 25 cm.

6.5. Casesandboxes
Cases of various design are used for root examinations. Wooden cases were
first used by Goedewaagen (1932, 1933), following Rotmistroff (1908) and
Maschhaupt (1915).The dimensions are 60x20x100 cm (Fig.59).Onewall
measuring 100x60cmis fastened by screws.The cases are filled with soil
and then buried with their surfaces level with the surrounding soil.Acrop
can be grown on the 60x20 cm top surface. When the crop is to be exami-

^ V ^ t ^ ^ ^ ^ ^ H H I P ^ P ^ P l H H H Fi §- 59 Wooden cases used

&?*-.s /ïV^Hl^lïlliÉtifi&afeSÖHI for root studies

62
Fig.60 Replacingonewallofawoodencasebyapinboard

ned, the wall fastened with screws is unscrewed and replaced by a pinboard
of the same dimensions. The pins are driven into the profile (Fig. 60).The
wholeis then turned upside down with the pinboard underneath and the
case is removed. The result is apinboard sample that can be washed and
further processed by the method described in Section2.3.
There are two types of concrete vessel in use. One type is a ready-made
vessel with dimensions 50x50x50cm, the walls being fastened together.
The onlyway to study root development in such boxesis byborings, as it is
hardly possible to obtain pinboard samples.
The second type is made of loose rectangular concrete slabs fitted together
to form abottomless container measuring l x l x l metre.It is advisable to
bury the slabs.After that the box can be filled with experimental soil.For
sampling ahole canbe dugon one side of the box, after which the adjoining
side-plate can be removed. After partly slicing off the soil to avoid marginal
effects, a pinboard sample can be taken. Eventually a second pinboard
sample canbe taken or the data can be completed by auger cores.
Aparticular method of root study is to observe root growfh behind awin-
dow-pane. Goedewaagen (1955) carried out experiments in small wooden
boxes with one vertical wall replaced by aglass panel. The inside areaof
these boxesis 10x10cm and the height 25 cm.Anetwork of squares was
arranged on the glass panel to facilitate observation. The boxes were slight-
ly tilted so that the glass wall leans forward. The glass wall should be

63
Fig.61 Smallwoodencaseswithoneglasspanel,providedwithsquares

covered during the experiment and the cover only removed when observa-
tions are made. These boxes are suitable for short term experiments in
which the root growth can be observed through the glass (Fig. 61).At the
end of the experiment the roots may bewashed free by removing the glass
panel and replacing it with a pinboard of suitable size.

6.6. Pot cultures

Different variations are possible. Use is made of ordinary flower pots,
Mitscherlich pots, and combinations of pots and water cultures. The glazed
earthenware vessels may also be mentioned under this heading as owing to
their size and shape theybelonghere rather than to the boxes.The pots can

64
 be filled in the same way as the cylinders. In the case of flower pots it
should be remembered that the diameter is not the same at all points.
Pot culture experiments are particularly useful when the plants are to be
examined at an early stage of growth and fullygrown root systems are not
required.
The disadvantages of pots are usually the shallow depth and slight volume,
that limit the root depth. Consequently the root systems of plants grown in
pots are often divergent from those grown under natural conditions.This is
further exaggerated bythe extremely intense root growth along the walls,
asis often seenin porous flower pots. Pot culture experiments are therefore
of limited use for root studies.This drawback is less applicable to the
glazed earthenware vessels which are 25 cmlong, 25 cmwide and 50cm
deep.
Roots of plants grown in pots can onlybe properly examined if the entire
root system iswashed free. The soil should be washed away in a downward
direction. Sometimes it is also possible to remove the entire contents from
the pot and then wash them.
Maschhaupt (1911) used a combination of flower pot and glass cylinder,
widening as far as possible the opening in the bottom of ordinary flower
pots so as to give the roots abetter chance of growing out of the pot. This
openingwas covered with a thin film of cottonwool to prevent soil from
falling into the beaker of nutrient solution underneath the pot. In the
middle of the pot a tapering wooden stick was placed with a diameter about
equal to that of the opening in the bottom. The soilwas pressed in round
the stick. After the stick had been removed the hole was filled up fairly
loosely with earth. Acavity was made in the middle in which a germinated
seed was placed. Thepots were placed in round holes made in cases with
detachable side walls, so that the beakers with nutrient solution were in the
dark. The root growth could be observed at regular intervals by removing
one of the side walls of the case.
Goedewaagen (1955) used a method which is substantially the same as
Maschhaupt's. Asbestos pots were used of which the bottom consisted of
wire netting with a2mm mesh.Thiswire netting was covered with a thin
layer of glass wool.The pots were filled with soil and placed on glass
cylinders filled with water or nutrient solution. In this way the subsoil was
imitated. Glasswool prevents soil particles from falling through the gauze
but permits free passage of the roots.The soil and water were separated by
a thin layer of air.
At the transition from pot to cylinder a metal collar was used to prevent
aeration and evaporation of the water in the cylinder, and to create a moist

65
 >• (i>: \i;*\'

lllll
O C' * C: i-i Fig.62 Viewofapot-cylinder
experiment

medium in the layer of air between the soil and the water. Adouble layerof
white polyethylene material was wrapped round the asbestos pots to pre-
vent evaporation through the cylinder wall (Fig.62).An experimental plan
of this kind is very suitable for studying the importance of subsoil roots for
the plant's supply of moisture. It is also possible to study fertilization pro-
blems in the topsoil and subsoil.
The soilin the pots was covered with alayer of gravel.The amount of water
consumed by the plants was calculated byweighing the pots and cylinders
separately at regular intervals.
The roots in the glass cylinders may be studied without making any further
provisions, and it is even possible to measure the longitudinal growth of the
roots during the experiment. The topsoil roots should be washed free.

6.7. Nutrient solutions

Zijlstra (1922) germinated seeds on hollow glass rings having a diameter of
13cm onwhich was stretched 1mm gauge gauze.Theringfloated in a jar
of water so that the gauze just touched the surface and the seeds placed on
it came into contact with the water. After germination the roots grew
through the meshes of the gauze and were freely suspended in the water.
Themain root was immediately marked. When it had attained a lengthof
some cm the cultures were transferred to nutrient solutions in 2V? litre
glass cylinders 24 cm high.
Water cultures are comparatively rarely used for root studies as conditions
differ so extensively from field conditions that it is difficult to compare the
conclusions. Ingeneral theymay be used for fertilizer and aeration pro-
blems. Cylinders or glassvessels of different sizes may be used for the
purpose. The growth of algae in the water or nutrient solution may cause
undesirable changes.This can be prevented by covering the vessel with
black paper, black polyethylene sheeting or corrugated paper. It may be
necessary to replenish and aerate the water regularly. The root system can
be easily studied by temporarily removing the cover.

7. Tracers

Wiersum has carried out some work with tracers and tracer-techniques. In
his publication (1967) he describes tracer techniques. He distinguishes
between application of tracers to the soil and to the plant and gives an
account of the amount of information obtained in both cases.

67
Appendices

Appendix 1 Schemefor description ofroot systems

1. Generalpicture
a. shape (e.g.square,oblong,etc.)
b. maximum depth incm
c. extentincm
d. colour
e. number of zonesdistinguished accordingto changeinthe shape of the
root system or in the density of the root zone

2. Descriptionbyzones
a. depthsincm
b. extentincm
c. density of rooting (in terms of sparse to abundant), e.g. < 2% sparse;
2-20% common; 20-70% many; 70-100% abundant
d. horizontal distribution of roots overthe width of theboard in thesoil
(in terms of regular or patchy)
e. colour

Appendix2 Nomenclature of roots

1. Monocotylodons
1.1 Annuals
1.1.1 From seed (cereals)
Main seminal root
Secondary seminal root branch roots androot hairs
Nodal roots
1.1.2 From bulbs or tubers (tulip, gladiolus, iris)
Adventitious roots
-Branch roots
-Root hairs
1.2 Perennials (pasture grasses)
Adventitious roots
-Branch roots
-Roothairs
(seedling axis,rhizome)

68
2. Dicotyledons
2.1 From seed (sugar beet, clover, lucerne)
Main root
- Branch roots
-Root hairs
2.2 From tubers (potatoes)
Adventitious roots
-Branch roots
-Roothairs
Adventitiousroots atthenodes
-Branch roots
-Root hairs
(Rhizomes, stolons are stems not roots)

Appendix3 Schemefor description ofindividual roots

I. Dicotyledons

1. Mainroot
a. shape (e.g.taproot, filiform, etc.)
b. lengthincm
c. thicknessinmm (wherenecessary at different depths or distances from
the base)
d. colour (colour mayvary accordingto soil composition)
e. number of zones,accordingto number ofprimarybranch roots
f. number of primary branch roots in each zone per unit length of the
main root.
g. length of the root-hair zone

2. Primary branchroots (perzone)

a. shape (e.g.elongated, twisted, etc.)
b. length incm(average + extremes)
c. thickness in mm (average+ extremes, wherenecessary at the base and at
various distances from the base)
d. colour '
e. number of zones,accordingto number of secondarybranch roots
f. number of secondarybranchroots,if necessaryperzone,interms-of few
to abundant perunit length
g. length of theroot-hair zone
3. Secondary branchroots
a. estimated length (average + extremes)
b. presence of tertiarybranch roots and of higher orders and their length
c. length of the root-hair zone

4. Adventitious roots (other roots)

details asunder1.1,1.2,1.3

5. Rootnodules
a. habit (single or multiple)
b. number
c. place

II. Monocotyledons (grasses and cereals)

1. Seminalroots
a. shape (e.g.elongated, twisted, etc.)
b. length incm (average + extremes)
c. thicknessinmm (average + extremes)
d. colour
e. number of zones,according tonumber of primary branch roots
f. number of primarybranch roots perunit length ineachzone

2. Primary branchroots
a. shape (elongated, twisted etc.)
b. lengthin cm (average + extremes)
c. thicknessinmm (average + extremes)
d. colour
e. number ofzones, according to number of secondarybranch roots
f. number of secondary branch rootsif necessaryperzoneinterms of few
to abundant per unit length

3. Secondary branchroots
a. estimated length of 2nd order branch roots
b. presence of tertiary branch roots and of higher orders andtheir length
c. length of the root-hair zone

4. Nodalroots
asII.l, II.2andII.3

70
Bibliography

1. Albrecht,D., 1951.Verbesserung der Spatendiagnose. Dt. Landwirt.

2:41-43.
2. Albrecht, D.,1955.Verbessertes Verfahren derFreilegung der Wurzel-
krone unter Beibehaltung ihrer natürlichen Lage.Arch. Gartenb.3:11-16.
3. Albrecht, D.,K.H.Fritzsche &C.Winkler, 1953.Weitere Entwicklung
des Strukturbohres. Dt.Landwirt.4:206-208.
4. Amson,F.W.van, 1954.Wortelonderzoek op enkele grasproefvelden van
het Lelydorpplan. Surin.Landb.2:229-235.
5. Anonymous, n.d. Root observation laboratory. Exp.Stn. S.Afr.Sugar
Assoc, n.p., 11 p.
6. Atherton Lee,H., 1926.The distribution of the roots of young D1135
plant cane in the soil.Hawaiian Planters Ree.30:267,511,520,523.
7. Bachy, A., 1964.Tropisme racinaire du palmier à huile. Oléagineux
19:684-685.
8. Bär, K.&O.Tseretheli, 1943.Der Einfluss der Schnitthäufigkeit auf die
Wurzelenwicklung junger Luzerne.Pflanzenbau 19:317-328.
9. Basset, D.M.,J.R.Stockton &W.L.Dickens, 1970.Root growthof
cotton asmeasured byP32uptake.Agron.J. 62:200-203.
10. Baumann, H. &M.L.Klaus, 1955.Ueber dieWurzelbildung bei hohem
Grundwasserstand. Z.Acker- u. Pflbau 99:410-426.
11. Bergann, F., 1964.Eineneuartige Trägerssubstanz zur erdelosen
Pflanzenkultur. Z.PflErnähr.Düng.Bodenk. 104:130-131.
12. Bhar, D. S., G.F.Mason &R.J.Hilton, 1969.Microscope trolley for
rhizotron. CanJ.Pl.Sci. 49:104-106.
13. Blaauw,A.H., 1938.Debeteekenis van den grondwaterstand voor de
bloembollencultuur. Verh.K.Ned.Akad.Wet. Afd. Natuurk. 2e Sectie,
371:91p.
14. Bloodworth, M.E., C.A.Burleson &W.R. Cowley, 1958.Root distribu-
tion of some irrigated crops usingundisrupted soil cores.Agron.J.
50:317-320.
15. Boehle,Jr. J., W.H. Mitchell, C.B.Kresge, &L.T.Kardds,1963.
Apparatus for taking soil-root cores.Agron.J. 55:208-209.
16. Bohne, H. &J. Garvert, 1951.Untersuchungen über die Bedeutung der
Ernterückstände des Getreides für die Humusversorgung. Z.PflErnähr.
Düng.Bodenk.55:170-178.

71
 17. Bonzon,B.,1964.Description et mode d'utilisation d'un appareil de
mesure photo-électrique des surfaces vegetables. Fruits (Paris)
19:577-581.
18. Bonzon, B.,&D.Picard, 1969.Matériel et méthodes pour l'étude dela
croissance et du développement en plein terre des systèmes racinaires.
Cah. ORSTOM,biol. 9:3-18.
19. Boonstra, A.E.H. R., 1931.Root systems of seven varieties of peas
grown under similar cultural conditions.Meded.LandbHogesch. Wage-
ningen 35,Verh.2:62p.
20. Borchert, H„ 1961.Ein methodischer Beitrag zur Entnahme von Boden-
proben in ungestörter Lagerung. Z.PflErnähr. Düng. Bodenk. 93:210-214.
21. Bosch, D.van den, 1952.Wortelonderzoek met eenvoudige hulpmidde-
len. Maandbl. Landbouwvoorlichtingsdienst9:335-341.
22. Bray, R.A.,J.B.Hacker &D.E.Byth, 1969.Root mapping of three
tropical pasture species using 32P. Aust.J.exp.Agric.anim.Husb. 9:445-448.
23. Brenchley, W.E.&V.G.Jackson, 1921. Root development in barley and
wheat under different conditions of growth. Ann.Bot. 35:533-556.
24. Brouwer, R., 1962.Influence of temperature of the root medium on the
growth of seedlings of various crop plants. Jaarb. Inst. biol. scheik.
Onderz. LandbGewass. 1962:11-18.
25. Brown, L.N., 1963.Thelacquer cement method of making soil mono-
liths. Bull.Calif, agric.Exp.Stn. 795:27p.
26. Buchele, W.T., 1961.Apower sampler of undisturbed soils.Trans.Am.
Soc. agric.Engrs4:185-191.
27. Bunt, A.C.&Z.J. Kulwiec, 1970.The effect of container porosity on
root environment and plant growth. I. Temperature. Pi.Soil 32:65-80.
28. Butijn, J., 1955.Bewortelingsproblemen in defruitteelt. In: De planten-
wortel in de landbouw. Min. Lanb.&Visserij, 's-Gravenhage :156-168.
29. Butijn, J., 1961.Bodembehandeling in de fruitteelt. Versl. Iandbouwk.
Onderz. 66.7:403p.
30. Cahoon, G.A.&E.S.Morton, 1961.An apparatus for the quantitative
separation of plant roots from soil.Proc.Am. Soc.hort. Sei.78:593-596.
31. Carley, H. E.&R.D.Watson, 1966.Anew gravimetric method for
estimating root-surface areas. Soil Sei.102:289-291.
32. Cerny, V., 1969. Metody morfologického a kvantitativniho studia
podzemnich orgânu rostlin. Studijni Informace 2:44p.
33. Clayton, M.F.&J.A.Lamberton, 1964.Astudy of root exudates by the
fog-box technique. Aust.J. biol. Sei.17:855-866.
34. Commonwealth Bureau of Pastures and Field Crops.Techniques for the
study of roots, 1960-1967.Bibliography S.1037, 6 p.

72
35. Coster, C, 1932.Wortelstudiën in de tropen. Meded. BosbProefstn.
Buitenzorg 31:96p.
36. Crist, J.W.&J. E.Weaver, 1924.Absorption of nutrients from subsoil
in relation to crop yield.Bot. Gaz.77:121-148.
37. Davies, D.B.&E.C.A.Runge, 1969.Comparison of rooting in soil
parent materials using undisturbed soil cores.Agron. J.61:518-521.
38. Dean, A.L., 1929.Root observation boxes.Phytopathology 19:407-412.
39. Doll, C. C, 1961.Apple tree root development in terraced loess soils.
Proc.Am.Soc.hort. Sei.78:1-7.
40. Eaton, F.M., 1941.Water uptake and root growth asinfluenced by
inequalities in the concentration of the substrate. Plant Physiol., Lancas-
ter 16:545-564.
41. Ellern, S.J., 1968.Are-usable experimental container for root studies.
Israel J. agric. Res.18:135-136.
42. Engel,K.H. &A.Raeuber, 1960.Einneues Kulturgefäsz zur laufenden
Beobachtung unterirdisch wachsender Organe (Kartoffeln). Züchter
30:206-207.
43. Fehrenbacher, J. B.&J. D.Alexander, 1955.Amethod for studying corn
root distribution using a soilcore sampling machine and shaker-type
washer. Agron.J.47:468-472.
44. Fergedal, L., 1967.Enmetod för framtvättning och undersökning av
rotsystem. Grundförbättring 20:53-60.
45. Flocker, W.J. &H. Timm, 1969.Plant growth and root distribution in
layered sand columns. Agron. J. 61:530-534.
46. Fox,R. L.&R. C.Lipps, 1955.Influence of soil-profile characteristics
upon the distribution of roots and nodules of alfalfa and sweet clover.
Agron. J. 47:361-367.
47. Frank, B., 1893.DieAssimilation des freien Stickstoffs durch die Pflan-
zenwelt. Bot. Ztg. 51:139-156.
48. Frankena, H.J.&M.A.J. Goedewaagen, 1942.Een vakkenproef over de
invloed van verschillende waterstanden op de grasgroeibij drie grond-
soorten.Versl.landbouwk. Onderz.48A:407-462.
49. Freckmann, W., 1926.Bericht über dieTätigkeit des Instituts für Melio-
rationswesen und Moorkultur. Landw.Jb. 64,Ergänzungsband1-11:34-62.
50. Frese,H.,W. Czeratzky &H.J.Altemüller, 1955.Ueber diéWirkung der
Verteilung organischer und anorganischer Düngerstoffe im Boden auf
Wurzelwachstum von Zuckerrüben. Z.PflErnähr. Düng.Bodenk.
69:198-205.
51. Fribourg,H.A., 1953.Arapid method for washing roots.Agron.J.
45:334-335.

73
 52. Gates, C.T., 1951.The quantitative recovery of root systems in pot
experiments. J.Aust. Inst, agric. Sei.17:152-153.
53. Gericke, S., 1946.DieBedeutung der Ernterückstände für den Humus-
haushalt des Bodens.II.Teil.DieErgebnisse eigener Untersuchungen.
Z.PflErnähr. Düng.Bodenk. 37:46-61.
54. Ghilarov, M.S., V.A.Kovda, L.N.Novichkova-Ivanova, L.E.Rodin,
V.M.Sveshnikova (Ed),1969.Methods of productivity studies in root
systems and rhizosphere organisms. Nauka, Leningrad, 240 p.
55. Gile, P.L.&J. 0. Carrero, 1917.Absorption of nutrients as affected by
the number of roots supplied with the nutrient. J. agric.Res. 9:73-95.
56. Gist, G.R.&R.M.Smith, 1948.Root development of several common
forage grasses to a depth of eighteen inches.J.Am.Soc.Agron.
40:1036-1042.
57. Gliemeroth, G., 1952.Wasserhaushalt des Bodens in Abhängigkeit von
der Wurzelausbildung einiger Kulturpflanzen. Z.Acker- u. PflBau.
95:21-46.
58. Gliemeroth, G., 1957.Untersuchungen über Ausbildung und Leistung
derKeim- und Kronenwurzeln beiSommergetreide. Z.Acker-u. PflBau.
103:1-21.
. 59. Goedewaagen, M.A.J., 1932.De groei van het wortelstelsel der planten
bij gelijke en bij ongelijke vruchtbaarheid van boven- en ondergrond.
Versl.landbouwk. Onderz.38A:179-199.
60. Goedewaagen, M.A.J., 1933.Het wortelstelsel der graanplanten bij
gelijke en ongelijke verdeling der meststoffen in de grond. Versl.land-
bouwk. Onderz.39A:243-266.
61. Goedewaagen, M.A.J., 1942. Het wortelstelsel der landbouwgewassen.
Departement van Landbouw en Visserij, Directie van de Landbouw,
's-Gravenhage, 173p.
62. Goedewaagen, M.A.J., 1948.Demethoden, die aan het Landbouwproef-
station en Bodemkundig Instituut T.N.O.te Groningen bij het wortel-
onderzoek opbouw- en grasland in gebruik zijn. Inst. Bodemvruchtbaar-
heid, Groningen, 11 p.
63. Goedewaagen, M.A.J., 1949.Een en ander over demethodiek van het
wortelonderzoek opbouw- en grasland. Maandbl. Rijkslandbouwvoorlich-
tingsdienst 6:194-200.
64. Goedewaagen, M.A.J., 1954.Beoordelingvan devitaliteit van het
wortelstelsel. Landbouwk. Tijdschr. 66:553-554.
65. Gooderham, P.T., 1969.Asimple method for the extraction and pre-
servation of an undisturbed root system from a soil. Pi. Soil 31:201-204.
66. Görbing, J., 1947.Die Spatendiagnose. Sachse, Hamburg, 32p.

74
67. Görbing, J., 1948.Die Grundlage der Gareim praktischen Ackerbau.
Landbuch Verlag, Hannover, 2 Bänder.
68. Gordienko, M., 1930.Ueber die Beziehungen zwischen Bodenbeschaffen-
heit und Wurzelgestaltung bei jungen Pflanzen. Landw.Jb. 72:125-139.
69. Haas, H.J., 1958.Effect of fertilizers, age of stand and decomposition
on weight of grass roots and of grass and alfalfa on soil nitrogen and
carbon. Agron.J.50:5-9.
70. Haas, H.J. &G.A.Rogier, 1953.Atechnique of photographing grass
roots in situ. Agron.J.45:173.
71. Hainnaux, G., 1966. Comparaison de deux méthodes de prélèvements de
racines sous prairie. ORSTOM, s.L, 50p.
72. Hall, N.S.,W.F.Chandler, C.H.M.van Bavel, P.H. Reid &J.H.
Anderson, 1953.A tracer technique to measure growth and activityof
plant root systems. Carol, agric. Exp.Stn.Tech. Bull. 101:40p.
73. Hanson, W.R.&L.A.Stoddart, 1940.Effects of grazing upon bunch
wheat grass.J.Am. Soc.Agron. 32:278-289.
74. Hanus, H., 1962.Wurzelprofil und Wasserversorgung der Grasnarbe bei
verschiedenen Grundwasserständen. Dissertation, Bonn, 74p.
75. Hayden, C.W.&W.H. Heinemann, 1968.Ahand-operated undisturbed
core sampler. Soil Sei. 106:153-156.
76. Hendrickson, A.H. &F.J. Veihmeyer, 1931.Influence of dry soil on
root extension. Plant Physiology, Lancaster 6:567-576.
77. Hesse,H., 1903.Beiträge zur Morphologie und Biologie der Wurzel-
haare. Inaug. Dissertation, Jena.
78. Hilton, R.J., D. S.Bhar &F.T.Mason, 1969.Arhizotron for in situ
root growth studies. Can. J. Pi. Sei.49:101-104.
79. Hoffmann, G., 1966.Verlauf derTiefendurchwurzelung und Feinwurzel-
bildung bei einigen Baumarten. Arch. Forstwesen (D.D.R.) 15:825-856.
80. Hopkins, H., 1953.Root development of grasses on revegetated land.
J. Range Mgmt 6:382-392.
81. Hudig, J., 1939.Klinisch grondonderzoek in de tropen. Landbouwk.
Tijdschr. 51:372-389.
82. Jacobs, E., D.Atsmon &U.Kafkafi, 1970.Aconvenient method of
placing radioactive substances in soil for studies of root development.
Agron. J. 62:303-304. '
83. Jacques,W.A., 1937.Anew type of root sampler. N.Z.J1Sei.Technol.
19:267-270.
84. Jacques, W.A., 1937.The effect of different rates of defoliation on the
root development of certain grasses.N.Z.J1Sei.Technol. 19:441-450.

75
85. Jager, A.&W.J.M.van der Voort, 1966.Collection and preservation of
soilmonoliths. Soil Survey Papers 2:24p.
86. Jiminez, E„ J. J. Salazar &V.Picado, 1967.Un método para medir el
volumen radical de plantas de cacao cultivadas en hydroponia. Turrialba
17:98-101.
87. Jonker, J. J., 1958.Bewortelingsonderzoek en ondergrondbewerking in
deNoordoostpolder. Dissertation, Wageningen, 164p.
88. Jutras, P.J. &A.C.Tarjan, 1964.A novel hydraulic soil auger-screen
shaker unit for the collection of root samples. Proc. Soil Crop Sei.Soc.
Fla.24:154-158.
89. Kampe, K., 1929. Studien über Bewurzelungsstärke und Wurzeleindrin-
gungsvermögen verschiedener Kulturpflanzen. Arch. PflBau 2:1-48.
90. Kauter, A., 1933.Beiträge zur Kenntnis des Wurzelwachstums der
Gräser. Ber.schweiz. bot. Ges.42:37-108.
91. Kawatake, M., G.Nishimura, K. Shimura &R. Ishida, 1964.Asoil-core
sampling machine and a fluctuating washer. Bull. Tokai-Kinki natn.agric.
Exp. Stn. 11:66-70.
92. Kelley, O.J., J.A.Hardman &D.S.Jennings, 1947.Asoil sampling
machine for obtainig two, three and four-inch diameter cores of un-
disturbed soil to a depth of sixfeet. Proc. Soil Sei. Soc.Am.12:85-87.
93. King, F.H., 1892, 1893.Natural distribution of roots in field soils.
A.Rep.Wis.agric. Exp.Stn. 9:112-120; 10:160-166.
94. Klapp,E., 1943.Ueber dieWurzelverbreitung der Gasnarbe beiverschie-
dener Nutzungsweise und Pflanzengesellschaft. Pflanzenbau 19:221-236.
95. Kmoch, H. G., 1952.Ueber den Umfang und einige Gesetzmäszigkeiten
der Wurzelmassenbildung unter Grasnarben. Z.Acker- u. PflBau
85:363-380.
96. Kmoch, H. G., 1957.DieWurzelarbeiten J.E.Weavers und seiner
Schule.Z.Acker- u. PflBau 104:275-288.
97. Kmoch, H. G., 1960.Die Herstellung von Wurzelprofilen mit Hilfe des
UTAH-Erdbohrers und ihre Ausdeutung. 1.Mitteilung. Methodik von
Wurzelprofil-Herstellung. Z.Acker- u. PflBau 110:249-254.
98. Kmoch, H. G. &H. Hanus, 1967.Die Herstellung von Wurzelprofilen
mit Hilfe desUTAH-Erdbohrers und ihre Ausdeutung. 4.Mitteilung. Zur
Veränderlichkeit der Durchwurzelung unter dem Einflusz von Boden und
Witterung. Z.Acker- u.PflBau 126:1-18.
99. Köhnlein, J. &H.Vetter, 1953.Ernterückstände und Wurzelbild. Paul
Parey, Berlin, 138p.
100. Kolesnikow, W.A., 1960.Metodika laboratornych i poljewych zanjatij

76
 poizutsjenioe kornewoi sistemy plodowych ijagodnych rastenij.Mosk.
Ord. Lenina sel'.-khoz. Akad. im K.A.Tïmirjazeva, Moskow, 58p.
101. Kolesnikow, W.A., 1962.Untersuchungen über das Wurzelsystem und
die Ernährung der Obstgehölze. Arch. Gartenb. 6/7:447-459.
102. Kolesnikow, W.A., 1962.Untersuchungen über Gesetzmäszigkeiten am
Wurzelsystem bei Obstgehölzen mittels einer besonderen Methode der
Probenentnahme. Tagber. dt. Akad. LandWiss. Berl. 35:169-177.
103. Könekamp,A., 1953.Teilergebnisse von Wurzeluntersuchungen.
Z. PflErnähr. Düng. Bodenk. 60:113-124.
104. Kulenkamp, A. 1969.Methodik der Untersuchung von Apfelwurzeln.
LantbrHögsk. Annlr. 35:1031-1039.
105. Kulescha, M., 1931.Oriënterend onderzoek over de ontwikkeling van
het wortelstelsel bij 2878 P.O.J,in verband met de grondsoort. Meded.
Proefstn. Java Suikerind. 8:317-359.
106. Kullmann, A., 1954.ZurAnfertigung und Bedeutung von Bodenlack-
filmen. Dt. Landwirt. 5:589-592.
107. Kullmann, A., 1957.Zur Intensität der Bodendurchwurzelung. Z.Acker-
u.PflBau 103:189-197.
108. Kutschera, L., 1960.Wurzelatlas mitteleuropäischer Ackerunkräuter
und Kulturpflanzen. DLG-Verlag,Frankfurt am Main.
109. Kvarazkhelia, T.K., 1931.Beiträge zur Biologie des Wurzelsystemes
der Obstbäume. Gartenbauwissenschaft 4:239-241.
110. Kvech, O., 1963. Prispevek k obahocoväni pudy poskliznovymi
zbytky plodin. I.Vojteskotravni smeska. Sb.vys.Sk.zemed. Praze
1963:193-201.
111. Lamba, P. S., H. L.Ahlgren &R.J. Muckenhirn, 1949.Root growth of
alfalfa, medium red clover, brome grass and timothy under various soil
conditions. Agron. J.41:451-458.
112. Lavy,T.L.&J.D. Eastin, 1969.Effect of soil depth and plant age on
32
Phosphorus uptake by corn and sorghmru Agron.J. 61:677-680.
113. Leonard, E.R.&G. C.Head, 1958.Technique and preliminary observa-
tions on growth of the roots of glasshouse tomatoes in relation to that
of the tops. J. hort. Sei.33:171-185.
• 114. Lieshout, J. W.van, 1956.De beworteling van een aantal landbouw-
gewassen. Versl.landbouwk. Onderz. 62:46p. f
115. Lieshout, J.W.van, 1961.Invloed van het bodemmilieu op ontwikke-
ling en activiteit van het wortelstelsel. Dissertation, Wageningen,91p.
116. Linkola, K.&A.Tiirikka, 1936.Ueber Wurzelsysteme und Wurzel-
ausbreitung der Wiesenpflanzen auf verschiedenen Wiesenstandorten.
Ann. Bot. Soc.zool.-bot. fenn. Vanamo 6,No. 6:207p.

77
 117. Lipps, R. C, R. L.Fox &F.E.Koehler, 1957. Characterizing root
activity of alfalfa by radioactive tracer techniques. Soil Sei.84:195-204.
118. Loewenstein, H., 1965.Aninjection probe for rapid placement of
radio-isotopes in soil.Proc. Soil Sei. Soc.Am.29:328-329.
119. Long, O.EL,1959.Root studies on some farm crops in Tennessee,
Bull.Tenn. agric. Exp.Stn 301:41p.
120. Low,A.J., 1951.The use of isotopes in agricultural research. II.
Chemy Ind. 1951:1124-1128.
121. Lyford, W.H. &B.F.Wilson, 1966.Controlled growth of forest tree
roots:Technique and application. Harv.Forest Pap. 16:12p.
122. Marsh, F.W.&L.T.Leonard, 1928.An apparatus for the superficial
examination of roots and nodules. Soil Sei.26:403-405.
123. Maschhaupt, J. G., 1911.Reactieverandering van de bodem ten gevolge
van plantengroei en bemesting. Versl.landbouwk. Onderz.10:50-93.
124. Maschhaupt, J. G., 1915.De beworteling onzer cultuurgewassen. Versl.
landbouwk. Onderz. 16:76-89.
125. Melhuish, F.M., 1967.Aprecise technique for measurement of roots
and root distribution in soils.Ann.Bot.32:15-22.
126. Melhuish, F.M.&A.R. G.Lang, 1968.Quantitative studies of roots in
soil. I. Length and diameters of cotton roots on a clay-loam soil by
analysis of surface-ground blocks of resin-impregnated soil. Soil Sei.
106:16-22.
127. Mesker, G.&J.J. Schuurman, 1972.Determination of root weights in
soil cores bymechanised and standardised freezing, in-vacuo peptisation
and washing. In press.
128. Metsävainio,K., 1931.Untersuchungen über das Wurzelsystem der
Moorpflanzen. Ann. Bot. Soc.zool.-bot. fenn. Vanamo 1,No. 1:418 p.
129. Murdock, J.T.&L.E.Engelbert, 1958.The importance of subsoil
phosphorus to corn. Proc. Soil Sei. Soc.Am.22:53-57.
130. Nad'jaranyi, F.M., 1939. Some data on the study of root systems and
plant mixtures. Sowet. Agron. 5:49-57.
131. Najmr, S., 1957.Ueber die Methode der Isolierung der Wurzelsysteme
der Futterpflanzen aus dem Boden. Z.Acker- u.PflBau 104:103-109.
132. Nelson, W.W. &R. R.Allmaras, 1969.Animproved monolith method
for excavating and describing roots. Agron.J. 61:751-754.
133. Newman, E.J., 1966.Amethod of estimating the total length of root
in a sample. J. appl. Ecol. 3:139-145.
134. Nightingale, H. J. &R.L.Smith, 1966.Aculturing unit to evaluate
sodic influence on calcium nutrition of alfalfa. Proc. Soil Sei.Soc.Am.
30:470-472.

78
135. Nutman,F.J., 1934.Theroot system of coffea arabica. Emp.J.exp.
Agric.2:293-302.
136. Obermayer, E., 1939.Root study technique. Herb. Rev.7:175.
137. Oskamp,J.&L.P.Batjer, 1933.Soilsin relation to fruit growingin
NewYork.III.Somephysical and chemical properties of the soils of the
Hilton and Morton Areas,Monroe County and their relation to orchard
performance. Bull.Cornell Univ.agric.Exp.Stn 575:1-34.
138. Osvald, H., 1919.Untersuchungen über die Einwirkung des Grund-
wasserstands auf die Bewurzelung von Wiesenpflanzen auf Moorböden.
Fühlings Landw. Ztg. 68:321, 370.
139. Otto, G., 1962.Ein Quotient zur Bestimmung der Verzweigungsdichte
von Gehölzfaserwurzeln im Zusammenhang mit Problemen derRhizo-
sphäre.Tagber. dt. Akad. LandwWiss. Berlin 35:179-184.
140. Pavlychenko, T.K„ 1937.Quantitative study of the entire root systems
of weed and crop plants under field conditions.Ecology 18:62-79.
141. Pavlychenko, T.K., 1937.The soil-block washing method in quantita-
tive root study. Can.J.Res. 15:33-57.
142. Pearson, R.W. &Z.F.Lund, 1968.Direct observation of cotton root
growth under field conditions. Agron. J.60:442-443.
143. Picard, D., 1969. Comparaison de deux techniques de prélèvement
d'échantillons de racines. Cah. ORSTOM,biol.9:19-31.
144. Pinkas, L.L.H., M.R.Teel &D.Schwarzendruber, 1964.Amethod for
measuring the volume of small root systems. Agron.J. 56:90-91.
145. Pinthus, M.J. & Y.Eshel, 1962. Observations on the development of
the root system of somewheat varieties. Israel J. agric.Res.12:13-20.
146. Price,K.R., 1965.A field method for studying root systems. Hlth Phys.
11:1521-1525.
147. Racz, G.J., D.A.Rennie &W.L.Hutcheon, 1964.The P32 injection
method for studying the rootsystem of wheat. Can. J. Soil.Sei.
44:100-108.
148. Raper, C.D.&S.A.Barber, 1970.Rooting systems of soybeans.
I.Differences in root morphology among varieties.Agron. J. 62:581-584.
149. Rappaport, J., 1938. Onderzoekingen over de ontwikkeling der wortels
bij Loliumpenene L.Meded. LandbHogesch. Gent 6:120-204.
150. Reicosky, D. C, R.J. Millington &D.B.Peters, 1970.Acomparison of
three methods for estimating root length. Agron. J. 62:451-453.
151. Reymerink, A., 1964.Eenverbeterde methode voor bewortelings onder-
zoek. Meded. Dir.Tuinb.27:42-49.
152. Richardson, S.D., 1956.Photosynthesis and root growth in tree
seedlings. Landbouwk. Tijdschr. 68:775-783.

79
 153. Riddler, A.M.H. &A.E.Thomas, 1966.Apower driven thin walled
core sampler. Agric. Gaz.N.S.W.77:627-632.
154. Rogers,W. S., 1969.TheEast Mailing root-observation laboratories.
In:W.J.Whittington (Ed.),Root Growth. Proceedings of the fifteenth
Easter School in Agricultural Science,University of Nottingham, 1968.
Butterworths, London, p.361-376.
155. Rogers,W. S.&M.C.Vyvyan, 1928.The root systems of some ten
year old apple trees on two different rootstocks, and their relation to tree
performance. A.Rep.E.Mailing Res.Stn Kent, II Suppl.14-15:31-43.
156. Roo,H. C.de, 1957.Root growth in Connecticut tobacco soils.Bull.
Conn, agric.Exp.Stn 608:36p.
157. Roo,H. C.de &L.K.Wiersum, 1963.Root training by plastic tubes.
Agron.J.55:402-405.
158. Rotmistroff, W. G., 1907.Die Gebiete der Verbreitung der Wurzeln
bei einjährigen Kulturpflanzen. Russisches Journal für experimentelle
Landwirtschaft 8: 530.
159. Rotmistroff, W. G., 1908.Die Gebiete der Verbreitung der Wurzeln bei
einjährigen Kulturpflanzen. Fortsetzung. Russisches Journal für experi-
mentelle Landwirtschaft 9:1-26.
160. Runge, E.C.A.1965.Thehydraulic soil-coring machine.A new method
of sampling Illinois soils.Illinois Res.7:10-11.
161. Sachs, J., 1860.Bericht über die physiologische Thätigkeit an der
Versuchsstation in Tharandt. Landw. Versuchsstn. 2:167,219.
162. Salim, M.H., G.W.Todd &A.M.Schlehuber, 1965.Root development
of wheat, oats and barley under conditions of moisture stress.Agron. J.
57:603-607.
163. Schulz, W., 1958.EinigeBeobachtungen über kurzfristige Veränderun-
gen im Boden auf kleinstem Raum.Z.PflEmähr. Düng. Bodenk. 80(125):
66-70.
164. Schulze,B.,1911.WurzelatlasI.PaulParey,Berlin.
165. Schulze,B.,1914.Wurzelatlas ILPaulParey,Berlin.
166. Schuurman, J. J., 1965.Influence of soil density on root development
and growth of oats. Pi. Soil 22:352-374.
167. Schuurman, J.J.&J.J. H. deBoer, 1970.The developmental pattern of
roots and shoots of oats under favourable conditions. Neth. J. agric.Sei.
18:168-181.
168. Schuurman, J.J. &M.A.J. Goedewaagen, 1955.Anewmethod for the
simultaneous preservation of profiles and root systems.Pi.Soil
6:373-381.

80
 169. Schuurman, J. J.&L.Knot, 1957.Het schatten van hoeveelheden
wortels in voor wortelonderzoek genomen monsters. Versl. landbouwk.
Onderz. 63.14:31p.
170. Schwendiman, J.L.,A.L.Hafenrichter &A.G.Law, 1953.The produc-
tion of tops and roots by grass and sweet clover when grown in mixtures.
Agron.J.45:110-114.
171. Simon, W.&D.Eich, 1955.Probleme und Methoden der Wurzel-
untersuchungen (unter besonderer Berücksichtigung leichterer Böden).
Z.Acker- u. PflBau 100:179-198.
172. Singer, F.P.&R.J. Hutnik, 1965.Excavating roots with water pres-
sure. J. For. 63:37-38.
173. Sioussaram, D., 1968.Observations préliminaires sur l'enracinement
desbananiers dans les sols de la Station deNeufchâteau Guadeloupe.
Fruits d'Outre Mer 23:473-479.
174. Slavikovâ, J., 1968.Die Beschleunigung des Auswaschens der Wurzeln
bei kwantitativer Rhizologie. Rostl. Vyroba 14:991-992.
175. Slëskin, P., 1908.Werden Nitrate von den Wurzeln assimiliert oder
nicht? Russisches Journal für experimentelle Landwirtschaft 9:32.
176. Smith, A.J. &L.A.Mabbitt, 1953.The contribution to soil organic
matter and nitrogen from cereal residues and undersown crops. J. Soil
Sei.4:98-105.
177. Solberg, P., 1935.Untersuchungen über die Wurzelausscheidungen
verschiedener Pflanzen bei ihrer Kultur ohne und mit Zusatz von Nähr-
salzen. Landw. Jb.81:891-917.
178. Sprague, H. B.,1933.Root development of perennial grasses and its
relations to soil conditions. Soil Sei.36:189-209.
179. Stanford, G.&J. D. DeMent, 1957.Amethod for measuring short-
term nutrient absorption byplants: I.Phosphorus. Proc. Soil Sei. Soc.Am.
21:612-617.
180. Stankov,N.Z., 1960.Methods for determining the length of plant
roots. Fiziologiya Rast. 7:736-739.
181. Stoecheler, J.H. &W.A.Kluender, 1938.The hydraulic method of
excavating the root systems of plants. Ecology 19:355-369.
182. Stransky, J.J. &M.V.Bilan, 1964.Power cutter for large undisturbed
soil cores.Agron. J.56:363.
183. Stuckey, J. H., 1941.Seasonal growth of grass roots.Am.J. Bot.
28:486-491.
184. Sutton, R.F.,1969.Form and development of conifer root systems.
Tech. Commun. Commonw. For. Bur., Oxford 7:130p.

81
185. Tackett, J. L.,E.Burnett &D.W.Fryrear, 1965.Arapid procedure for
securing large, undisturbed soil cores.Proc. Soil Sei.Soc.Am.29:218-220.
186. Taerum, R.&M.D. Gwynne, 1969.Methods for studying grass roots
under East African conditions and some preliminary results. E.Afr. agric.
For.J. 35:55-65.
187. Tavcar, A.& V.Kendjelic, 1966.Characteristics of the roots of seed-
lings produced in linen rolls.Indicators of the growth of the plants under
field conditions. Arh.poljopr. Nauke 19:151-159.
188. Thames, J.L.&R.D.Reynolds, 1961.A hydraulic soil sampler. Agric.
Engng, St. Joseph, Mich. 42:431-432.
189. Tollenaar, D.,1930.Onderzoekingen over de ontwikkeling van het
wortelstelsel van tabak onder verschillende omstandigheden; tevens
bezien in verband met cultuurmaatregelen enbrandbaarheid. Meded.
Proefstn Vorstenlandsche Tabak 65:47p.
190. Troughton, A., 1957.The underground organs of herbage grasses.
Bull. Commonw. Bur. Pastures Fid Crops 44:163p.
191. Tucker, M.&C. von Seelhorst, 1898.Der Einfluss, welchen der Wasser-,
gehalt und der Reichtum des Bodens auf dieAusbildung derWurzeln und
der oberirdischen Organe der Haferpflanzen ausüben. J.Landw.64:52-63.
192. Ueno, M.,K.Yoshihara &T. Okada, 1967.Livingroot system distin-
guished by the use of Carbon-14.Nature, Lond. 213:531-532.
193. Upchurch, R.P., 1951.The use of trench-wash and soil-elution
methods for studying alfalfa roots.Agron. J.43:552-555.
194. Vetter, H. &S.Scharafat, 1964.DieWurzelverbreitung landwirtschaft-
licher Kulturpflanzen imUnterboden. Z.Acker- u. PflBau 120:275-298.
195. Voigt, E., 1949.DieAnwendung der Lackfilm-Methode bei der Bergung
geologischer und bodenkundlicher Profile. Mitt. geol. Staatsinst. Hamburg
19:111-129.
196. Voort, W.J.M.van der, 1970.Literature about taking and preservation
of soil monoliths. Soil Survey Inst. Wageningen, No. 4846, 9 p.
197. Weaver, J.E., 1926.Root development of field crops. McGraw-Hill,
New York, 291p.
198. Weaver, J. E.&R.W.Darland, 1947.Amethod of measuring vigor of
range grasses.Ecology28:146-161.
199. Weaver, J.E.&R.W.Darland, 1949.Soil-root relationships of certain
native grasses in various soil types.Ecol.Monogr. 19:303-338.
200. Weaver, J.E.,J. Kramer &M.Reed, 1924.Development of roots and
shoots of winter wheat under field environment. Ecology 5:26-50.

82
201. Weaver, J.E.&Ellen Zink, 1946.Length of life of roots of ten species
of perennial range and pasture grasses. Plant Physiol., Lancaster
21:201-217.
202. Welbank, P.J.&E.D.Williams, 1968.Root growth of a barley crop
estimated bysamplingwith portable powered soil-coring equipment.
J. appl.Ecol.5:477-481.
203. Weller,F.,1964.Vergleichende Untersuchungen über dieWurzelVertei-
lung von Obstbäumen in verschiedenen Böden des Neckarlandes.Arb.
Landw. Hochschule, Hohenheim 31:1-181.
204. Wetzel, M., 1958.Der Futterroggen, die Futterroggengemische und die
sonstige Futtergemische des Winterzwischenfutterbaues als Lieferanten
von Futter und Wurzeln. Wiss.Z.Univ.Rostock 7:89-366.
205. Whitcomb, C.E.,E.C.Roberts &R. Q.Landers, 1969.Aconnecting
pot technique for root competition investigations between woody plants
or between woody and herbaceous plants. Ecology 50:326-328.
206. Wiersum, L.K., 1958.Density of root branching as affected by sub-
strate and separate ions.Acta bot.neerl. 7:174-190.
207. Wiersum, L.K., 1967.Potential subsoil utilization by roots.PLSoil
27:383-400.
208. Wiersum, L.K., 1967.Root-system development. In: Soil-Moisture and
Irrigation Studies. International Atomic EnergyAgency, Vienna, p.83-95.
209. Wiklert, P., 1960.Studier av rotutvecklingen hos nâgra nyttoväxter
med särskild hänsyn tili markstrukturen. Grundförbättring 3:113-148.
210. Willard, C.J. &G.M.Mc Clure, 1932.The quantitative development
of tops and roots in blue grass with an improved method of obtaining
root yields.J.Am.Soc.Agron. 24:509-514.
211. Williams,F.E.&H. K.Baker, 1957.Studies on the root development
of herbage plants. I. Techniques of herbage root investigations. J. Br.
Grassld. Soc.12:49-55.
212. Williamson, R.E., 1968.Influence of gas mixtures on cell division and
root elongation. Agron. J. 60:317-321.
213. Witte, K., 1929.Beitrag zu den Grundlagen des Grasbaus. Landw.Jb.
69:253-310.
214. Woods,F.W., 1969.Root extension of forest trees: a method of
investigation using radioactive tracers. In: W.J.Whittington (Ed.),Root
Growth. Proceedings of the fifteenth Easter School in Agricultural
Science,,University of Nottingham, 1968.Butterworths, London, .
p.413-417.

83
215. Wijk, W. R.van, 1952.De bepaling van dewerkzaamheid der wortels
met behulp van radio-active isotopen. Landbouwk. Tijdschr. 64:398-399.
216. Yoder, R.E., 1936.Adirect method of aggregate analysis of soils and
a study of the physical nature of erosion losses. J. Am. Soc. Agron.
28:337-351.
217. Zillmann, K.H., 1956.Beobachtung desWurzelwachstums in Feld-
beständen. Dt. Landwirt. 7:394-400.
218. Zijlstra, K., 1922.De hoofdwortel van enige graansoorten. Versl.land-
bouwk. Onderz.26:19-59.

84
Key to the bibliography

I General:
32, 34, 54, 61, 62, 63, 96,100,101,102,108,184,190, 203, 211

II Field experiments:
a. auger: 1, 2, 3, 14, 15, 18, 20, 21,26, 37, 43, 56, 65, 69, 71, 73,74, 75,83,
84, 88, 91, 92, 97, 98, 130, 143, 148, 153, 156, 160,171,176,180,182,185,
186, 188, 194, 202, 203, 211,216
b. soil excavation: 16, 19, 23, 46, 51, 53, 66, 67, 80, 81,84, 93, 94, 95, 99,
103, 105, 110, 113, 116, 119, 131,140, 141,145, 170, 178, 183, 193, 199,204,
209, 210
c. pinboard: 4,18, 21,44, 81, 87,114,124,128,132,148,149,156,168,186,
204
d. profile wall investigation:
- washing: 21,23, 70, 73,151,181,193
- mechanical means (e.g. needle): 7, 21,28, 29,104,105,121,137,156,173,
197, 209, 213
e. (spatial) position of entire root system:
-washing: 50, 90, 93,156,172,186
- mechanical means: 35, 39,104, 109,113,116, 121,128,135,155,189, 200
f. underground observation of roots in situ (glass panels):
5,12,13, 78, 79,113,136,142,154,158,164,165, 217
g. radioactive tracers: 9, 22, 72, 82,112,115,117,118,120,129, 146,147,
192, 208, 214, 215

III Container experiments

a.,pot cultures: 10, 23,27, 28,29,42, 47, 68,76, 77, 89, 90, 111, 149,179,
191, 198,205,206, 218
b. boxes with glass panels: 8, 38, 90,113,152,162,163,177
c. pot-plus cylinder combination: 123,134
d. cylinders: 10, 41,45,48, 58, 89,150,166,167, 200, 201,207
e. large containers: '
- pinboard: 59, 60,124,138,157,159
- washing: 6,19, 36, 49, 136
/. water cultures: 24, 40, 47, 55,149,161,175, 218
g. spray (mist) cultures: 33, 212
h. radioactive tracers: 9,192, 215

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IV Specialtechniques
a. washingmethods: 18,30,43,44,51,52,69,91,98,103,127,141,148,
174
b. thin-layersoilmonoliths: 25,85,106,168,195,196
c. surfacearea, lengthandvolume measurements: 17,31, 57,86,99,107,
126,133,135,139,144,150,180, 200
d. vitality andactivity: 64,192
e.estimations: 169
/. undisturbed soilcores ascultureunits: 3,37,74,182
g. specialaids:2,11,12, 42,65,70,89,102,121,122,125,151,187,201,
205,207

86