82 SYMPOSIUM II

FACTORS AFFECTING SOIL MOISTURE -

PLANT GROWTH RELATIONS1
ROBERT M. HAGAN -
Department of Irrigation, University of California. Davis, Ca!if., U.S.A.

SUMMARY
The gross effects of deficient and of excessive soil moisture on plant growth are well known, but controversy has existed
for many years around the question whether the so-called “'available moisture” is
equally available for plant growth or available only with such increasing difficulty that plant growth
functions are retarded before the wilting point is reached. Existing viewpoints are illustrated by
schematic diagrams and the probable effects of plant, soil, weather, and some other miscellaneous
factors on. soil rnoisture-plant growth relations, discussed.
Considered are the theory that plants can obtain water with equal facility between field capacity and
permanent wilting percentage; the notion that growth diminishes progressively as soil moisture falls
below field capacity; and the theory that plant functioning is directly related to the tenacity with
which water is held by soil. Also considered are views that physiological activity may be related to
time integrated soil moisture stress or to maximum soil moisture stress."
Various measurable aspects of plant 'growth' (elongation, increase in fresh or dry weight and vege-
tative versus reproductive development) do not respond in the same manner to increasing moisture
stress. Appreciable shifts may occur m the relative abundance of certain chemical constituents within
plants. Indicated are possibilities of so controlling soil moisture stress over the growing season as to
favour production of that plant organ or constituent for which the crop is grown. Greater emphasis on
such considerations would lessen confusion now existent in our understanding of plant-soil-water
relations and thus in irrigation recommendations.
Rooting characteristics of well established perennials and the expanding root systems of annual crops
are discussed. Where plants have few widely spaced roots, soil moisture determinations may give
quite a false picture of moisture conditions at root surfaces. Such crops will respond to irrigations,
although measured soil moisture stress is quite low.
Problems of relating plant growth to soil moisture stress conditions varying with both time and soil
depth are considered. The question is raised whether any type of integrated or average soil! moisture
stress value, which includes some low stress values from portions of the root zone containing few roots,
can represent the true stress experienced by plants. On the other hand, deep roots absorbing water
against low soil moisture stresses mask the effects of relatively high soil moisture stresses occurring
over parts of root systems. Some problems of studying plant responses to soi5 moisture conditions
in small containers are mentioned.
Soil factors affecting density or depth are reviewed and several ways weather factors influence soil,
-moisture-plant-growth relations summarized. When the sizable number of plant, soil, weather, and
miscellaneous factors influencing the relation of soil moisture to yield of some specific plant organ or
constituent, are considered, it is not surprising that conflicting results have been obtained even in
irrigation experiments involving some given crop. This discussion suggests the probable impossibility
of finding any one relation between crop yields and moisture depletion or soil moisture stress, at least
as measured with our present methods.
Present information, though meagre in many respects, allows some fairly accurate predictions whether.
1
Paper read by J. R. FURR.

Reprinted from the Report of the XIVth International Horticultural Congress, The Netherlands. 1955
Published by H. Veenman & Zonen, Wageningen (Holland)
R. M. HAGAN 83

growth or yield in given situations is likely to be unaffected by depletion of nearly all available water
or is likely to be increased by irrigation at low soil moisture stresses. Table IV lists those conditions lowering the
probability that yields will be increased by irrigation at relatively low soil moisture
stresses, and Table V conditions raising the probability that yields will be increased by avoiding
relatively high soil moisture stresses. Mention is made of some economic and other factors which
under practical conditions may of irrigations.

The gross effects of deficient and of excessive soil moisture on plant growth are, of course, well
known. With irrigation and, where necessary with drainage, the farmer has the opportunity to
exercise greater control over soil moisture than he does over any of the other soil physical
factors.
A lively controversy has existed or many years around the question, whether the so called
available moisture is equally available for plant growth, or available only with such increasing
difficulty that plant growth functions are retarded before the wilting
point is reached. These viewpoints have been summarized in recent review articles
by VEIHMEYER and HENDRICKSON [12] and by RICHARDS and WADLEIGH [5]. This
difference of opinion is more than academic, for our answer to the farmer's very
practical question, “when should I irrigate”, depends upon our understanding of soil
moisture-plant growth relations.
It is not particularly helpful for the present purpose to recite the evidence that supports
these two viewpoints. Instead, our efforts should be directed towards an understand-
ing of some factors which have contributed to this controversy. The now voluminous
literature on plant-soil-water relations includes many reports of irrigation experiments
on given crops from which seemingly diametrically opposed conclusions have been
drawn. This situation is most confusing, especially to those of you who must attempt
to apply the results of research to the practical problem of crop production. The im-
portant task now is to attempt to analyze carefully the conditions prevailing in the
various studies which have been reported, with the hope of learning why competent
investigators have been led to such divergent conclusions.
I shall attempt to illustrate by schematic diagrams the existing viewpoints and to
suggest the probable effects of plant, soil, water, and some other miscellaneous
factors, on soil-moisture-plant-growth relations. In attempting this, I am well aware
that much of what follows involves some speculation and may someday be shown to
have been quite erroneous. In assuming this risk, I take some comfort from a remark,
attributed to FRANCIS BACON, that “truth arises more easily from error than from
confusion”. The diagrams presents may contains some errors, but I hope they may
serve to lessen the confusion now prevalent.

PLANTGROWTH RESPONSES WITHIN ONE IRRIGATION CYCLE

Growth rate not diminished over available range
Let us consider first, plant growth responses as the available soil moisture is depleted
within one irrigation cycle. VEIHMEYER and HENDRICKSON's theory, that plants can
obtain a supply of water with equal facility between field capacity (FC) and the per-
manent wilting percentage (PWP), can be illustrated by figure 1. This represents their
view (VEIHMEYER and HENDRICKSON, [11, 13]) that rate of growth is not diminished
84 SYMPOSIUM II

over the available range or, in other words, that no measurable increases in rate of
growth are obtained by irrigating until the soil moisture falls to near the permanent
wilting percentage. The question mark indicates the uncertainty in the vicinity of
the permanent wilting percentage.

FIG. 1. Schematic
representation of
VEIHMEYER and
HENDRICKSON'S theory of
“equal availability”.

Growth rate progressively diminished as soil moisture content falls below FC

On the other hand, it has been maintained that plants respond favourably to relatively
high soil moisture conditions, and the statement is sometimes made that plant growth
diminishes progressively as the soil moisture content falls below field capacity and
ceases at the permanent wilting percentage. Such a picture, figure 2, of soil-moisture-
plant-growth relations is not uncommon and often creeps into textbooks and popular
articles, although, as Í understand it, there is little support for this idea among research
workers in this field. This may be called the “more water, more growth” idea.
'The foregoing notion arises in part from numerous experiments which have suggested
that plant growth is related to the tenacity with which the water is held by the soil.

Growth rate as a function of soil moisture stress

The 'availability' of soil moisture is now frequently described in terms of soil moisture
tension which is dependent upon surface forces, and in terms of total soil moisture
stress (SMS), which includes all surface and osmotic forces arising from the presence
of solutes in soil solution. The soil moisture-component-moisture-stress curves of figure 3
illustrate the moisture retention characteristics of four non-saline soils of different
textures (sands to clays). In saline soils, it is considered important to include the os-
motic effects. Moisture content-soil moisture stress relations for a given soil, to which
increasing amounts of soluble salt had been added, are given in figure 4.
The theory that plant growth is a function of soil moisture stress can be expressed
R. M. HAGAN 85

Fig. 2. Representation of view

that plant growth is directly
related to soil moisture content
within this available range.

O 25 50 75 100
AVAILABLE MOISTURE DEPLETION, PER CENT

Fig. 3. Idealized moisture

retention curves for a non-
saline sandy soil (curve 1),
loam soil (curve 2), and two
clay soils (curves 3 and 4)

O 25 50 75 100
AVAILABLE MOISTURE DEPLETION, PER CENT

Fig. 4. Relations between

soil moisture depletion and
(total) soil moisture stress
for Panoche loam to which
had been added sufficient
salt to give and osmotic
pressure of the soil solution
at approximately the field
capacity of nearly O
atmospheres(curve 1), 2
atmospheres (curve 2), and 4
atmospheres (curve 3).
(Redrawn from
WADLEIGH, 1946)

AVAILABLE MOISTURE DEPLETION. PER CENT

86 SYMPOSIUM II

FIG. 5. Representation of theory

that plant growth is a function of
soil moisture tension or total soil
moisture stress. For non-saline
sandy (curve 1) and clay (curve 2)
soils and for saline soil (curve 3)

100

AVAILABLE MOISTURE DEPLETION, PER CENT

diagrammatically as in figure 5. If such a relation exists, little retardation in growth

on the non-saline sandy soil would be expected until nearly all the available water
had been depleted (curve 1), but on the non-saline clay soil some slowing of growth
should occur after about 50 per cent. depletion (curve 2). On the saline soil (curve 3),
a somewhat reduced growth rate would be expected at moisture contents near field
capacity, and it would decline appreciably even in the upper half of the available
range.

CUMULATIVEGROWTH OVER SEVERAL IRRIGATION CYCLES

So far consideration has been confined to growth rates within one irrigation cycle. If
moisture is equally available for plant growth over the available range, then total
growth over a period of time will be independent of the level of moisture depletion
permitted before irrigations. This relation is indicated by curve 1 of figure 6. If, on

FIG. 6. Curve 1 represents

total growth over a series
of cycles for irrigations at
given levels of moisture
depletion as predicted by
the theory of equal
availability. Curve 2 gives
total growth as predicted
by the theory that growth is
a function of time
integrated soil moisture
stress. Curve 3 indicates
total growth on the basis
that brief periods of 100
high moisture stress may
have an exaggerated effect
on plant growth
R. M. HAGAN 87

the other hand. growth rates are related to soil moisture stress, then total growth over
a period during which. the stress varies may be related to some average stress condition.
WADLEIGH [18] proposed that cumulative growth be related to a moisture stress
function integrated (averaged) over time. Total growth over the time interval of one
irrigation cycle or of several cycles calculated on this basis is represented by curve
2 of this figure. Its shape will depend on that of the soil moisture stress-moisture
content curve. On the other hand, some field studies have suggested that yields are
related to maximum stress prevailing prior to irrigation. WADLEIGH and other
workers [1, 6] recognize that an integrated moisture stress may not adequately ex-
press the physiological effect of the extreme variation in moisture stress over the
irrigation cycle and suggested that high moisture stress values, though present íbi'
only a brief time interval, might have an exaggerated effect on plant response. This
possibility is depicted by curve 3. Further work will be needed to determine the im-
portance of high stress periods and the relative effects on growth of moisture stress and
specific ion toxicities in saline systems. Before leaving this figure, it should be pointed
out that much greater decreases in yield would occur, if all the available moisture is
depleted in the major portion of the root zone during one or more of the irrigation
cycles. Doubtless the serious reductions in yield, often attributed to treatments calling
for depletion of nearly all the available water, have been caused by failure to recognize
that actually all the available water had been exhausted over much of the root system.
The occurrence of wilting may not always be a reliable indicator of such moisture ex-
haustion, for under field conditions some crops do not wilt until long after growth has
been checked.

OTHER FACTORS INDEPENDENTLY AFFECTING PLANT GROWTH

Considerable experimental support can be found for each of the generalized soil
moisture - plant growth relations represented in figure 6. This situation indicates that
moisture-growth relations must be greatly influenced by the interplay of other factors
which independently affect plant growth. The possible effects of some plant, soil,
weather, and other miscellaneous factors, are discussed in the following sections.

Plant Factors
Insufficient attention to the specification of what is meant by plant growth has often
complicated the interpretation of the results of irrigation experiments. We easily
recognize several different manifestations of growth, such as elongation of plant organs,
increase in fresh or dry weight, and vegetative versus reproductive development.
These common-place processes are resultants of intricate combinations of many
physiological processes which are probably not all equally affected by increasing soil
moisture stress and an accompanying change in the internal water balance of cells
and tissues. Thus it is not surprising that various measurable aspects of growth do
not respond in the same manner to moisture stress. Greater emphasis on this point
would aid materially in lessening the confusion now existent in our understanding of
plant-soil-water relations and thus of irrigation needs.
88 SYMPOSIUM II

Available Moisture Depletion per cent

FIG. 7. Effect of moisture depletion on rate of petiole elongation during dark and light periods for ladino clover grown in
containers in a controlled environment room (hagan, et al, 1951)

FIG. 8. Effect of moisture depletion on green weight yields, dry weight yields, and dry matter percentage of forage from
ladino clover grown in containers (HAGAN, et al, 1951).
R. M. HAGAN 89

Fig. 9. Effect of
moisture depletion on
photosynthesis in ladino
clover grown in
containers under
continuous light and at
constant temperature
and humidity. (Adapted
from UPCHURCH et al,
1956)

A number of workers have shown that certain aspects of growth are more affected
by moisture stress than others. Data from recent studies on ladino clover (HAGAN
et al [3]) illustrate this point. In figure 7 the rate of petiole elongation for clover grown
in containers is shown to decrease markedly with rising soil moisture stress and during
the light periods to approach zero before wilting appears. In another container ex-
periment, figure 8, green weight production declined with increasing moisture stress,
while dry matter production was largely unaffected until at least 75 per cent of the
total available moisture had been depleted. This suggests that, although cellular elon-
gation may be retarded, the rates of photosynthesis and respiration must be little
affected over a considerable range of moisture stress. The net rate of carbon dioxide
exchange for clover, grown in containers under continuous light for two drying cycles,
are given in figure 9. This suggests that the rate of photosynthesis was not dimin-
ished appreciably until about the time wilting appeared. The respiration rate was
little affected over this stress range. These results are in general agreement with the
findings of others that photosynthesis and respiration are relatively insensitive to

FIG. 10. Differences in the

effects of soil moisture stress
on several aspects of growth:
Curve (1) photosynthesis or
respiration, curve (2) dry
weight yield, and curve (3)
elongation or fresh weight
yield
90 SYMPOSIUM II

moisture stress (for a discussion see: RICHARDS and WADLEIGH [5]). Differences in
the effects of soil moisture stress on several aspects of growth for one plant, ladino
clover, are summarized schematically in figure 10.
As plants are subjected to increasing moisture stress, appreciable shifts in the relative
abundance of a variety of chemical constituents may occur in some plants, see figure
11. The rubber content of guayule has been shown to rise under increased stress
(HUNTER and KELLEY [4]). The percentage of sugar in cane and beets can also be
raised by moisture stress (WIERSMA [15]). In tobacco, increasing stress is reported to
have lowered the sugar content and raised the percentage of nicotine and nitrogen in
the cured leaves. (VAN BAVEL [10]). Thus, the economic value of a crop may be in-
fluenced appreciably by moisture stress, particularly during the period of maturation.
Differences in the response of various growth processes to moisture stress conditions
point the way to the possibilities of so controlling the soil moisture stress over the
growing season as to favour the production of that constituent or plant organ for
which the crop is grown. The effects of given soil moisture stress conditions on the
yields of field corn have been reported to depend upon the stage of growth. Corn
appears to be particularly sensitive to moisture stress during the tasseling period.

AVAILABLE MOISTURE DEPLETION, PER CENT

FIG. 11. Rubber content of guayule as affected by irrigation within the available range and by de-
pletion of available moisture on Hesperia sandy loam. In treatment A the soil moisture was
maintained near field capacity at all times (29 irrigations), in treatment B an average moisture
depletion of 25 % was permitted in 12-24 inch depth (18 irrigations), and in treatment C an
average moisture depletion of 67% was permitted in the same depth of soil (11 irrigations).
Treatment D received one pre-irrigation in early April which wetted the soil to a depth of
over 8 feet. By harvest on August 8, nearly all the available water in treatment D was de-
pleted from the 8-fooí depth. Treatment E received no irrigations and by August was at or
below the PWP throughout the sampling depth. (Adapted from HUNTER and KELLEY, 1946)
R. M. HAGAN 91

FIG. 12. Difference in effects of moisture

depletion on dry weight yield of shrub
and on yield of rubber per acre.
(Adapted from HUNTER and KELLEY, 1946)

Vegetative vigour is not necessarily associated with a comparable degree of produc-

tivity. This is well illustrated by studies on guayule, figure 12, which show that in-
creased moisture stress retards the vegetative growth of the shrub, but leads to a higher
production of rubber.
Another plant factor of extreme importance in determining the relation between
measurable soil moisture stress and plant growth is the nature of the root system.
Different interpretations of root development and of moisture conditions within the
soil penetrated by roots contribute to the existence of contradictory views on plant-
soil-water relations. Under favourable soil and growing conditions, most perennial
crops develop well-branched root systems which thoroughly
permeate the soil depth characteristic of the plant. Below
this depth, the spatial density of absorbing roots diminishes
until so few remain that moisture extraction cannot be de-
tected. This depth distribution of roots is illustrated by the
drawing in figure 13 and leads to a moisture depletion pattern
of the type shown here for bermuda grass (figure 14). This
illustration calls attention to several features of a root system
of importance in analyzing water relations and raises ques-
tions about what is meant by the loosely used term “root
zone”. It is desirable to distinguish what may be called the
'complete extraction zone' from the 'partial extraction zone'
or 'transition zone'. The complete extraction zone may be
defined as the volume or, in the case of closely spaced plants,

FIG. 13. Fibrous root system of grass showing decrease in root density
with depth (Adapted from SPRAGUE, 1940)
92 SYMPOSIUM II

the depth from which all the available water has been removed when the crop ceases
growth. The soil within this zone is thoroughly permeated with fine roots. Here the
complete extraction zone would extend to a depth of about 2 feet. The transition
zone contains a varying concentration of roots usually decreasing with distance from
the plant and here extends to a depth of at least 6 feet. It should be remembered that
in isolated plants, especially when young or planted on a wide spacing, the root zone
is three-dimensional so that the transition zone may form a considerable volume
around the complete extraction zone. The existence of this transition zone often
seriously complicates the interpretation of moisture-growth experiments.

FIG. 14. Differences in moisture depletion

under bermuda-grass reflecting decreasing
numbers of absorbing roots with greater
depth. (Unpublished studies by HAGAN
and MADISON, Univ. of Calif., Davis)

Analysis of moisture conditions within the expanding root system of an annual

crop is even more complicated. In the seedling state, only a tap root or a few branch-
ed roots penetrate the soil. Some annuals rapidly develop a well-branched root
system which permeates an ever-enlarging soil volume. At the same time, some roots
grow out ahead into a correspondingly enlarging transition zone. Thus, if the soil
has been previously wet to field capacity through a considerable depth, these growing
roots continuously come into contact with additional supplies of available water at
low tensions. If the roots are well-branched and grow rapidly enough, they may
contact new supplies of readily available water with sufficient rapidity to meet the
needs of transpiration. A crop such as watermelon on the deep alluvial soil at Davis
does not respond to irrigation, although a relatively high soil moisture stress may
develop within an ever-increasing soil volume.
Other annuals send out a few widely spaced roots which leave large volumes of un-
explored soil between roots, particularly in the early stages of growth. Figure, 15 shows
the skeletal root system of a young sweet corn plant. Soil moisture samples or even
R. M. HAGAN 93

FIG. 15. Skeleton root system of young sweet com growing in pervious deep soil. (Unpublished
studies by DONEEN, Univ. of Calif., Davis)

moisture indicating devices may give quite a false picture of moisture conditions at
the root surface. Crops with sparse-roots will respond to irrigations although the
measured soil moisture stress may be quite low. The sparser the roots, the greater
the likelihood that growth will be retarded by delaying irrigation (figure 16). Very
similar varieties of beans have shown deferent responses to irrigations which are re-
lated to differences in their root development.1
Thus, the fraction of the available range which can be utilized before growth is checked
will vary with root density. Were it possible to sample only the soil adjacent to ab-
sorbing roots, we might find that the actual moisture stress conditions were consider-
ably higher than indicated by our present methods. Until methods are developed
to measure the moisture stress experienced by plants, it apparently will be necessary
in the case of sparsely-rooted crops to establish some rather arbitrary moisture deple-
tion limits, which unfortunately will depend on stage of growth, growing conditions,
and on such other factors as soil and weather.
It is not uncommon to find reports that crop yields declined, unless irrigated at some
given soil "moisture stress. Often it is difficult to learn at what depth the stress was
measured and seldom is its relation to rooting depth and spatial density indicated.
Where spatial density diminishes rapidly with depth, the soil moisture stress corre-
sponding to any plant response will vary markedly over a short depth interval.
1
Unpublished studies at the University of California, Davis, by D. W. HENDERSON and L. D. DONEEN.
94 SYMPOSIUM II

FIG 16. Schematic represen-

tation of the effect of spa-
RELATIVE RATE OF GROWTH, PER CENT

100
tial density of roots on the
relation of growth to the
apparent depletion of
available moisture

25 50 75 100
AVAILABLE MOISTURE DEPLETION, PER CENT

The suggestion has been offered (RICHARDS & WADLEIGH [5]) that roots in the
transition zone absorbing water against low soil moisture stresses may
mask the effects of relatively high soil moisture stresses occurring
over only a part of the root system. Thus, it may be argued that whenever the
transition zone extends through a considerable volume of soil, depletion of most of
the available moisture from the complete extraction zone may have relatively little
influence on growth. This idea is represented schematically in the diagram of figure
17. Whether the crop will respond to increasing stress within its complete extraction
zone, or will be unaffected, will depend on whether the roots in the transition zone can
supply water fast enough under the prevailing conditions to maintain the necessary
water balance in the plant. The presence of an unknown fraction of the total absorb-
ing roots in the transition zone and the continued elongation of the roots into moist

FIG. 17. Schematic

representation of the relation
between the portion of the
roots subjected to stress and
25 50 75 rate of growth
AVAILABLE MOISTURE DEPLETION, PER CENT
R. M. HAGAN 95

soil, particularly in annual crops, makes it very difficult to evaluate the actual soil
moisture stress to which the plant is subjected. Growth responses to increasing soil
moisture stress in container experiments may be much larger than experienced in
field experiments set up under as nearly comparable conditions as possible. This is
illustrated in figure 18 by the relative yields of ladino clover for comparable con-
tainer and field experiments. The smaller decrease in yield produced by increasing
soil moisture stress within the complete extraction zone in the field experiments may
be attributable to those roots in the transition zone which are in contact with moisture
at relatively low stress. For these and several other reasons, container experiments
may not be reliable indicators of crop response to soil moisture stress under field con-
ditions.

Soil Factors
Soil properties and conditions listed in table I have a decided effect on soil moisture -
plant growth relations.

Any soil factor which affects root density or depth can be expected to influence the
response of the crop to irrigation. Mechanical impedance, slow water penetration
and poor internal drainage, and deficient aeration, are frequently responsible for sparse
and shallow roots.
TABLE I. Soil factors affecting soil moisture-plant growth relations

SOIL STRUCTURE, TEXTURE AND DEPTH

Mechanical impedance
Infiltration rate
Internal drainage rate
Aeration
Moisture retention characteristics (soil moisture versus soil moisture stress relations)
Unsaturated hydraulic conductivity

WATER TABLE
SALINITY
Soil moisture stress
Toxicity
Soil structure

PLANT DISEASES AND NEMATODES

TEMPERATURE

FERTILITY

Soil structure, texture, and depth determine the total capacity of the soil for
storing available water for plant growth. The total available moisture capacity within
the root zone and the moisture-release characteristics of the soil are both important
factors determining the rate of change in soil moisture tension or stress. Deep-rooted
crops on deep soils usually show smaller responses to irrigations than shallower
rooted crops on the same soil. Crops growing on a soil in which 75 to 85 per cent. of
96 SYMPOSIUM II

FIG. 18. Relative green weight yields

for ladino clover under corresponding
irrigation treatments in containers
and in field plots. (Adapted from
HAGAN et al, 1951)

the available water is released at tensions below one atmosphere may be expected
to show a smaller response to irrigations at a given moisture depletion level than the
same crops growing on a soil in which less than 50 per cent. is released at such low
tensions. Great emphasis has been placed on the 'energetics' of moisture availability
in recent years, but doubtless the 'dynamics' must also be considered. The rate at
which water can move to the absorbing root surface may play an important role in
plant-soil-water relations.
A stable water table in the lower portion of the normal root zone of a crop may
supply considerable portion of the water absorbed by the roots and make the plants
less responsive to moisture changes in the soil above the capillary fringe. On the other
hand, a fluctuating water table may increase crop responses to early irrigation by
restricting live roots to a shallow depth. High water tables in semi-arid regions usually
lead to the accumulation of salts. As mentioned previously, salinity may affect soil
moisture-plant growth relations by decreasing moisture availability through increased
soil moisture stress, by interfering with root growth and absorption through toxicity
reactions, and by contributing to poor soil structure which, in turn, influences infiltra-
tion, drainage, aeration, and root growth.
Soil-borne plant diseases and nematodes, by reducing root surface, may cause
crops to respond favourably to irrigations at seemingly very low moisture stress levels.
Soil temperature also affects the rate of root growth and their distribution with depth
(HAGAN [2]).
The fertility status of the soil and possibly the depth distribution of some
essential element may also determine the growth response of crops to irrigations at
various moisture depletion levels. These data (Table II) illustrate the effect of nitrogen
applications on the response of corn to two moisture tension levels. Without nitrogen
R. M. HAGAN 97

fertilizer, the additional irrigations required to maintain low tensions lowered yields
presumably as a result of leaching out some nitrogen. The difference in yield between
the relatively wet and dry treatments increased with greater amounts of fertilizer. In
soils where the available supply of some essential element is confined to the top-soil,
drying out of the upper portion of the root zone may seriously retard plant growth,
even though the plant may still be adequately supplied with water. In a current ex-
perítnent1 with sugar beets growing on a phosphate fertilized soil, leaf analyses show
a phosphate deficiency whenever the surface soil dries out, although deeper roots are
obtaining sufficient water to maintain adequate plant turgor. In such cases yield in-
creases produced by more frequent irrigations are not chargeable directly to an inade-
quacy of moisture supply. Fertility responses have complicated the interpretation of
many soil moisture versus plant growth experiments.

TABLE II. Corn yield as affected by nitrogen and soil moisture

tensión*

Nitrogen Yield of corn (bushels/acre)

applied Low High Difference
(Ib/acre) SMT SMT low-high
0 62 70 -8
40 95 88 7
120 130 105 25
240 142 111 31
* Irrigated when soil moisture tension (SMT) at 9 inch depth in row reaches 0.8 atm. (Iow SMT)
and 4.1 atm. (high SMT). (Adapted from SINGLETON et al. [7]).

Weather Factors
Weather factors can influence soil moisture-plant growth relations in several ways, as
summarized in table III. Time permits little more than a mention of these factors.
Weather conditions, particularly light and temperature, may so influence the
growth characteristics of the shoot and root as to affect soil-moisture-growth relations.
Current work by DONEEN2 suggests that late-planted sugar beets, which must develop
roots during hot dry weather, fail to develop as dense or deep a root system as early
seeded beets. The length of the crop season before autumn rains or frost may at least
partially determine whether harvestable yields will be affected by imposing different
soil moisture stress levels during the growing period.
Meteorological factors-light, temperature, humidity, and wind-control
the rate of water loss by transpiration and evaporation. Where water use rates are
high, crops will deplete the available soil moisture more rapidly, and growth may be
more affected, by increasing soil moisture stress. Plant growth is probably dependent
upon plant turgor or plant moisture stress, whose relation to soil moisture stress for

1
Unpublished studies at the University of California, Davis, by HILLS et al, 1954.
2
Unpublished studies at the University of California, Davis, 1954.
7
98 SYMPOSIUM II

FIG. 19. Schematic represen-

tation of the possible effect
of transpiration rate on
soil moisture-plant
growth relations

25 50 75
AVAILABLE MOISTURE DEPLETION, PER CENT

different rates of transpiration needs to be explored. It can be reasoned that an in-

creased rate of transpiration would lower the plant turgor corresponding to any given
soil moisture stress and would have the effect of causing growth to diminish at higher
moisture levels. This notion is illustrated schematically in figure 19. Much more
work on this point is needed.

TABLE III. Weather factors affecting soil moisture — plant growth relations
GROWING CONDITIONS
Light
Temperature
Length of growing season

EVAPORATING CONDITIONS
Light
Temperature
Humidity
Wind

Miscellaneous Factors
Problems associated with insect control or harvesting may at times influence the
apparent effects of soil moisture conditions on crop yields. A study of forage and seed
production by ladino clover provides an interesting example. As shown in figure 20,
increased dryness reduced forage yields but increased the harvestable yield of seed.
The total seed actually produced by the clover also diminished with increased stress,
but the higher humidity associated with the wettest treatment caused such a serious
pre-harvest loss of newly produced seed as to reduce the harvestable yields at the end
of season below those on the drier treatments. Some type of harvesting problem may
affect the results of soil moisture-plant growth experiments more frequently than is
realized. .
R. M. HAGAN 99

AVAILABLE MOISTURE DEPLETION, PER CENT

FIG. 20. Difference in effects of moisture depletion on harvestable yields of vegetative and reproduc-
tive phases of growth in ladino clover. (Adapted from HAGAN et al, 1951)

Mention has been made of a sizable number of soil, plant, weather, "and other mis-
cellaneous factors, which may influence the effects of various moisture depletion levels
on plant growth or yield of some specific organ or constituent. When viewed against
this background, it is not at all surprising that conflicting results have been obtained
even in irrigation experiments involving some given crop. It also seems that this dis-
cussion emphasizes the probable impossibility of finding any one relation between
crop yields and soil moisture depletion or soil moisture stress, at least as measured with
our present methods. The problem is further complicated by the plant itself. There
is some evidence that growth is related to the turgor of plant tissues, but turgor ís not
a function of soil moisture stress alone. Rather it is dependent upon the relative rates
of water absorption and transpiration and upon the time interval over which any differ-
ences in rates may have continued. Until more is learned about the effects of turgor
on various physiological processes and about other factors which may influence soil
moisture-plant growth relations, our approach to problems of soil moisture control
must continue to involve considerable empiricism.
Present information, meagre though it is in many respects, may allow us to make some
fairly accurate predictions whether growth or yield in given situations is likely to be
unaffected by depletion of nearly all the available water as measured by our present
methods, or is likely to be increased by irrigation at lower soil moisture stresses. At
the very real risk of some error, two check lists have been prepared in the following
two tables. Listed in the first (Table IV) are conditions which may lower the probability
that yields will be increased by irrigations at relatively low soil moisture stresses. The
100 SYMPOSIUM II
other (Table V) lists conditions which raise the probability that yields will be increased
by avoiding relatively complete depletion of the available soil moisture. These are
merely lists, and it is not implied that all conditions must be present. Relative weight-
ing of each of these conditions is not considered. If a given situation is described by
some entries from both tables, prediction will be much more difficult.

TABLE IV. Conditions which lower the probability that crop yields are increased by irrigations at
low soil moisture stress

PLANT
Deep, dense, fast-growing roots
Xerophytic characteristics
Dry weight yields of reproductive organ desired
Harvest for content of sugar, oil, etc.

SOIL
Deep soil; good structure
Good infiltration, internal drainage, aeration
Large fraction of available water held at low tensions
Non-saline
Fertility level low; nutrients distributed in profile
Constant water table in reach of roots

WEATHER
Planted well ahead of hot dry weather
Major growth period before hot dry weather
Low evaporation rates

TABLE V. Conditions which raise the probability that crop yields are increased by irrigations at
low soil moisture stress

PLANT
Shallow, sparse, slow-growing roots
Fresh weight yield of vegetative organ desired
Quality dependent upon size of vegetative organ

SOIL
Shallow soil; poor structure impeding root growth
Slow infiltration and intemal drainage; poor aeration
Root disease, nematodes present
Small fraction of available water held at low tensions
Saline soils or water
Fertility level high; nutrients concentrated in top-soil
Very high soil temperatures, with shallow rooted crops

WEATHER
Planting at beginning of hot dry weather
Major growth period during hot dry weather
High evaporation rates

Up to this point, we have been concerned only with the physiological response of
plants to soil moisture. No mention has been made of economic and other
R. M. HAGAN 101

factors which under practical conditions may dictate the timing of irrigations. It is often
questionable whether the increased yields, sometimes obtainable under rela-
tively frequent irrigations, will pay for the added cost of water and labour. Even those,
opposed to the concept of equal availability as a scientific theory, are in general agree-
ment that the following practical considerations all suggest the desirability of a sparing
use of irrigation water: maximum use of limited water supplies, water and nutrient
losses caused by deep percolation, danger of developing a drainage problem through
over-irrigation, problems of soil structure, quality of the marketable product, and
problems of plant disease and longevity in perennial crops, which are often aggravated
by frequent irrigation. Although these considerations are of real importance in deter-
mining farm irrigation practices, their relative importance differs from place to place
and even from year to year. Much confusion would be avoided in preparing irrigation
recommendations, if the relative importance attached to soil moisture-plant growth
responses and to other considerations was clearly indicated.

FIG. 21. Irrigation before

complete moisture deple-
tion is generally necessary
under actual irrigation con-
ditions in order to ensure
a continuous supply of
available moisture. This
practice also avoids the
portion of the available
range where the largest
differences in growth re-
sponse would be predicted
by the different theories on
plant-soil-water relations

One other practical problem should be mentioned. The 3 curves of an earlier figure
(figure 6) represent the considerable differences in yield to be predicted on the basis of
existing theories. One point, however, should be mentioned. In order to ensure a
continuous supply of available soil moisture and to allow for unforeseen delays in irri-
gation or unusually dry weather, the irrigation farmer generally cannot plan to allow
nearly complete depletion of the soil moisture; but, to allow for a margin of safety,
he should plan to irrigate while some available moisture still remains. The fraction
of the total available moisture range which can be utilized with safety depends on a
number of factors including crop rooting characteristics, the soil and the irrigation
system. In figure 21, let us consider the effects of allowing a safety margin of 15 per
cent. If such an allowance is made to meet the practical problems of irrigation under
farming conditions, then a considerable portion of the differences predicted by these
several theories on plant-soil-water relations tends to disappear. However, to raise
102 SYMPOSIUM II

irrigation efficiency and to increase crop production, it is highly important that vigorous
programs of research be directed towards increasing our knowledge of soil moisture-
plant growth relations.

REFERENCES
[1] BERNSTEIN, L., and PEARSON, G. A., 1954. The influence of integrated moisture stress achieved
by varying the osmotic pressure of culture solutions on the growth of tomato and pepper plants.
Soil Sci. 77, 355-368.
[2] HAGAN, R. M., 1952. Soil temperature and plant growth. In: Agronomy II. Soil physical con-
ditions and plant growth, B. T. Shaw, editor. Academic Press, New York, pp. 367-447.
[3] ———, PETERSON, M. L., UPCHURCH, R. P., and JONES L. G., 1951. Relationships of soil.
moisture stress to different aspects of growth in ladino clover. (Manuscript submitted to Proc.
Amer. Soil Sci. Soc.).
[4] HUNTER, A. S., and KELLEY, O. J., 1946. The growth and rubber content of guayule as affected
by variations in soil moisture stresses. J. Amer. Soc. Agron. 38, 118-134.
[5] RICHARDS, L. A., and WADLEIGH, C. H., 1952. Soil water and plant growth. In: Agronomy U.
Soil physical conditions and plant growth, B. T. Shaw, editor. Academic Press, Inc., New
York. pp.73-251.
[6] ROBINS, J. S., and DOMINGO, C. E., 1953. Soma effects of severe soil moisture deficits at
specific growth stages of corn. Agron. J., 45, 618-621.
[7] SINGLETON, H. P., et al., 1950. Soil, water and crop management investigations in the
Columbia Basin Project. Washington Agr. Exp. Sta. Bul. 520, 29-30.
[8] SPRAGUE, HOWARD B., 1949. Better lawns for homes and parks. McGraw-Hill, Inc., New York.
[9] UPCHURCH, R. P., PETERSON, M. L., and HAGAN, R. M. The effect of soil-moisture content on
the rate of photosynthesis and respiration in ladino clover (Trifolium repens L.) Plant Phys. 30,
297-303.
[10] VAN BAVEL C. H. M., 1953. Chemical composition of tobacco leaves as affected by soil
moisture conditions. Agron. J. 45, 611-614.
[11] VEIHMEYER, F. J., and HENDRICKSON, A. H., 1950(a). Essentials of irrigation and cultivation of
orchards. Univ. of Calif. Agr. Ext. Service Circ. 50, 23 pp.
[12] ——— and ———, 1950(b). Soil moisture in relation to plant growth. Ann. Rev. Plant
Physiol. 1,285-304.
[13] ——— and ————, 1952. The effects of soil moisture on deciduous fruit trees. Report 13th
Int. Hort. Congress, 1, 306-319.
[14] WADLEIGH, C. H., 1946. The integrated soil moisture stress upon a root system in a large con-
tainer of saline soil. Soil Sci. 61, 225-238.
[15] WIERSMA, D., 1955. Soil moisture conditions and sugar accumulation in the sugar beet. Ph. D.
Thesis, University of California.