Root growth and soil-water extraction by winter and

spring wheat
M. H. Entzl, K. G. Gross2, and D. B. Fowler
Crop Development Centre, University of Saskatchewan, Saskatoon, Saskatchewan, Canada

Can. J. Plant Sci. Downloaded from pubs.aic.ca by 50.26.13.38 on 10/24/14

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S7N 0W0. Received 24 Dec. 1991, accepted 20 May 1992.

Entz, M. H., Gross, K. G. and Fowler, D. B. 1992. Root growth and soil-water extraction by winter
and spring wheat. Can. J. Plant Sci. 72: 1109-1120. Efficiencies ofsoil-water extraction and use
are important factors determining crop productivity in dryland regions. This study was conducted from
1986 to i988 to compare root growth and soil-water extraction of Norstar and Norwin winter wheat
(WW) (T riticum aestivum L.) and Katepwa and HY320 spring wheat (SW). Total root length and rooting
depth increased with crop-development stage and level ofavailable soil water. Preanthesis root development was concentrated at soils depths with temperatures greater than 15"C. Maximum rooting depth
at crop maturity ranged from 1 10 to 130 cm and 60-80% of the total root length was found in the
top 50 cm of soil. The rooting patterns of SW and WW were similar at each sampling date despite
differences in the stages of development. However, the earlier development of WW resulted in a greater
root length for WW by the end of May, an advantage maintained until WW maturity. Differences in
the rooting patterns of semidwarf and tall cultivars were not detected. The average change in available
soil water (0-130 cm depth) over all cultivars and locations between early May and crop maturity was
5 cm. Ninety-five percent of this water was extracted from the upper 70 cm of soil. Actual rooting
depth and maximum depth of soil-water extraction were highly correlated (r : 0.87**). In six of eight
trials, WW cultivars used significantly more water from the top 50 cm of the profile in May than SW.
In one trial, WW extracted significantly more water from several soil-depth increments than SW over
the entire growing season. In the remaining trials, levels of available soil water at SW anthesis and
harvest were similar for al1 cultivars or were higher for WW cultivars because of soil-water recharge
by late-season precipitation. Therefore, with the exception of greater early-season water extraction by
WW, f'ew consistent differences in soil-water extraction Datterns were observed amons cultivars
considered in this study. Higher water-use efficiencl ior WW in these trials was attributed to diiferences
in timing of water extraction and not to differences in depth of water extraction.
Key words: Canadian prairies, soil-water extraction, root growth, spring wheat, winter wheat, semidwarf,

tall, Triticum

aestivLtm

Entz, M. H., Gross, K. G. et Fowler, D. B. 1992. Croissance racinaire et extraction de I'eau du

sol par le bl6 d'hiver ou de printemps. Can. J. Plant Sci. 72: 1109- I 120. L'efficacit6 de I'extraction
et de l'utilisation de 1'eau du sol est un d6terminant important de la productivit6 des cultures dans les
zones de culture sdche. Une 6tude rdalis6e de 1986 ir 1988 avait pour objet de comparer la croissance
racinaire et I'extraction de I'eau du sol chez les bl6s d'hiver (BH). Norstar et Norwin et les b16s de
printemps (BP) Katepwa et HY320 (Triticum aestirum L.). La longueur totale des racines et la profondeur
d'enracinement augmentaient en fonction du stade de d6veloppement de la plante et du niveau d'eau
disponible dans le sol. Le d6veloppement racinaire en pr6-anthdse se concentrait dans la couche du
so1 poss6dant une temp6rature supdrieure zl 15'C. A maturit6, la profondeur d'enracinement maximale
allait de 110 A 130 cm, et 60-80% de la longueur totale des raciens 6tait comprise dans les 50 premiers
centimdtres du sol. Le mode d'enracinement de BP et de BH 6tait le m6me iL chaque date de pr6ldvement,
malgr6 Ies diff6rences du stade de croissance des divers cultivars. Toutefois, le d6veloppement plus
lPresent address: Department
of Plant Science, University of Manitoba, Winnipeg Manitoba, Canada R3T 2N2.
rPresent address: Ducks Unlimited, Box 28, Shoal Lake, Manitoba, Canada ROJ 120.

Can. J. Plant Sci.72: 1109-1120 (Oct. 1992)

l 109

Can. J. Plant Sci. Downloaded from pubs.aic.ca by 50.26.13.38 on 10/24/14

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1110

CANADIAN JOURNAL OF PLANT SCIENCE

pr6coce de BH lui a valu une plus grande longueur de racines h la fin de mai, avantage qu'il a conservd
jusqu')r maturit6. On n'a pas observ6 de diff6rences d'enracinement entre les cultivars demi-nains et
les cultivars ir paille longue. Le changement moyen dans la teneur en eau du sol disponible (0-130 cm)
entre le d6but de mai et la maturit6, tous cultivars et emplacements confondus 6tait de 5 cm. Quatrevingt-quinze pour cent de cette eau etait tir6 des 70 premiers centimbtres du sol. La profondeur d'enracinement rdelle et la profondeur maximale d'extraction de l'eau du sol 6taient fortement corr6l6es (r : 0,87x+).
Dans six essais sur huit, les cultivars de BH utilisaient significativement plus d'eau des 50 premiers
centimdtres du sol en mai que les cultivars BP. En outre, ils soutiraient significativement plus d'eau
que BP de plusieurs couches de profondeur durant l'ensemble de la saison de v6g6tation. Dans les autres
essais, les niveaux d'eau du sol disponible d I'anthdse 6taient les m0mes chez tous les cultivars, ou encore
ils 6taient plus elev6s chez les BH h cause de la r6alimentation de la nappe par les pr6cipitations de
fin de saison. Il semble donc qu')r part la plus forte extraction de I'eau du sol par les BH, il y avait
peu de diff6rences r6gulibres du mode d'extraction parmi les cultivars examin6s. La plus forte efficacit6
d'utilisation de I'eau observ6e chez les BH est expliqu6e par des diff6rences dans le calendrier d'extraction
de I'eau durant la saison et non par des diff6rences de la profondeur d'extractron.

Mots cl6s: Prairies canadiennes, extraction de l'eau du sol, croissance racinaire, bl6 de printemps, bl6

d'hiver. demi-nain.

)r

paille lonsue. Triticum aestiyum

Maximum root-penetration and soil-waterextraction depths reported for wheat range

from 70 to 200 cm (Hurd 1968; Lupton et al.
1974:'Black et al. 1981). However, in most
studies the majority of water extraction and
root growth of wheat has been limited to the
upper 70 or 90 cm (Brown 1971; Doorenbos
and Kassam 1979 Proffrtt et al. 1985).
Depth of rooting and soil-water extraction
of wheat depends on both environmental and

genetic factors. Schurmann (1959)

and

Proffitt et al. (1985) reported that when water

supply in the upper soil profile was plentiful,
rooting depth remained shallow. Sharma and

Ghildyal (1977) observed that drier soil

conditions promoted root elongation and
increased the degree of branching, allowing

the crop to exploit soil water more

completely. However, Barraclough et al.
(1989) reported that wheat crops do not neces-

sarily exhaust all available soil water before

commencing uptake from the next deeper
layer. Normally, root growth in wheat is most
rapid between tillering and anthesis (Conor
1975). However, availability of water can
strongly modifu this pattern, resulting in high
growth rates even after anthesis (Campbell
et a\. l91l; Barraclough et al. 1989).
Lupton and Bingham (1970) suggested that

semidwarf wheats have longer and more

extensive root systems than older, tall types.

However, subsequent studies (Lupton et al.

19'74; Cholick et al. 1977) found no significant correlation between plant height and
rooting depth or soil-water-extraction patterns
among winter wheat cultivars. Black et al.

(1981) reported that winter wheat (WW)

rooted deeper than spring wheat (SW);
however, Evans and Wardlaw (1976) cited
evidence that the growth of a root system is
dependent on the length of the vegetative
period and not growth habit.
There are many different techniques for
measuring rooting patterns in crops. Bohm
(1979) concluded that the profile-wall method
was an accurate and quantitative method that
was less time-consuming than traditional core
methods for the direct collection of root data.
Measurements of soil-water depletion have
been used to determine rooting patterns
indirectly (Davis et al. 1965; Bond et al.
1971; Campbell et al. 19'71; Proffitt et al.
1985; Bauer et aJ. 1989).
Winter wheat is a relatively new crop to
most of the Canadian prairie region. To date,

little is known about root growth and soilwater extraction of WW compared with tradi-

tional SW cultivars. Consequently, the first

objective of this study was to compare root
growth and soil-water extraction of tall and
semidwarf WW and SW cultivars across a
wide range of environments in the semiarid

ENTZ ET AL.

WHEAT ROOT GROWTH AND SOIL-WATER EXTRACTION

and dry subhumid regions of Saskatchewan.

Water-use efficiency (WUE) has been shown
to be significantly higher for WW than SW
in western Canada (Entz and Fowler 1991).
Therefore, an additional objective in this
study was to determine if the WUE advantage of WW was related to differences in root
growth and total soil-water extraction of WW

included in the Clair 1988 trial. Norstar, Katepwa

and the semidwarf cultivars Norwin winter wheat

and HY320 spring wheat were included in the

remaining trials. Weather, cultural methods,

fertilization, placement of neutron access tubes,
evapotranspiration (ET), phenological development, grain yield, and water-use efficiency for
these studies have been reported in a related study

(Entz and Fowler 1991). Experimental design for

trial was a randomized complete block with
three or four replicates. Plot width was 2.5 m in
all trials. Plot length was 30 m in one of four replicates in each trial that direct root measurements

and SW cultivars.

each

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MATERIALS AND METHODS

Direct root observations were made in trials

at

in

1985 (Elstow clay loam; Canada Soil

Survey Committee 1978; previous crop, chemical
summerfallow of wheat stubble), Floral in 1986
(Bradwell clay loam; previous crop, canola), Outlook in 1986 (Asquith fine sandy loam; previous
crop, canola), and Clair in 1986 (Yorkton loam;
previous crop, barley). Soil-water extraction was
monitored to provide an indirect measure of the
pattern of root development in these trials and in
additional trials conducted at Hagen (Blaine Lake
loam; previous crop canola), Paddockwood (Pelly
loam; previous crop canola), and Clair in 1987
(previous crop barley) and Clair in 1988 (previous
crop barley). The tall winter wheat cultivar Norstar
was evaluated in the Perdue 1985 trial. Norstar and
the tall spring wheat cultivar Katepwa were
Perdue

1111

were made. Plot length was 3 or 5 m in the

remaining replicates of all trials.
The profile-wall method (Bohm 1979) was used
to obtain a direct measure of root growth in all 1985
and 1986 trials. Measurements were made on one
replicate per site. Sampling was conducted at four
different crop-development stages (Table 1). The
procedure was as follows: A backhoe was used to
cut a trench perpendicular to the plots. The trench
was approximately 2 m wide by 2 m deep and long

enough to include all plots in a replicate. The

vertical wall corresponding to each plot was
smoothed with a flat-bottomed spade and then a

profile knife (Bohm 1979). We exposed the roots

by removing a soil layer approximately 1.3 cm
thick from the vertical face. This was accomolished

Table l. Percentages of total root length at different depths and total root length (m roots m-r soil) in the soil
profile for winter (Norstar and Norwin) and spring (Katepwa and HY320) wheat grown at Perdue in 1985 (Norstar
only) and Clair, Floral and Outlook, Saskatchewan, in 1986; the profile-wall method was used to obtain estimates
oI root length
1986

Perdue 1985
Soil

19 June

3 July

depth

ZGS 65

15

(cm)

Norstarz

Norstar

Date

21

sw

IJ

ww
%

0-10

t-1

l0-30

25

30-50
50-70
70-90

at
10

90-t l0

664
21
28
20

29
1

Date 2

Date 3

35

65

SW

ww

65

sw

89

Date 4
90

ww

sw

of total root length

60
26
145

64
30

17
2

61 48
31 23
691317
29816
1

40
25

30

1l

l9

110-130

Total root
rengtn (m

SE

m ')

218

129

24.9

zZadoks etal. (1974) growth

stage: 21, start oftillering; 31-35, early stem elongation; 65, anthesis; 75, milk;

89 90. mature.

vDate l, 26 May to 3 June; date 2, 16 to 26 June; date 3, 10 to 17 July; date 4, 30 July to

SW, spring wheat; WW, winter wheat; SE, standard error of the means.

l8

August.

r112

CANADIAN JOURNAL OF PLANT SCIENCE

by spraying the trench face with water

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under
pressure (0.276 MPa) from a teejet nozzle on a
hand-gun attached to a water pump. The wash
pattern (up and down and then from side to side)
and time (3-6 min, depending on soil type) was
similar for all plots within a location in 1986. At
the Perdue site, wash time per plot was greater for
the 3 July sampling date than the 19 June one.
Trenches used to sample roots at different growth
stages within the same replicate were at least
3 m apart.
A 35-mm slide (photograph) was taken of the

vertical face

of the profile wall to provide

soil-water extraction (Lupton et al. l9'74; Cholick

et al. 1977; Bauer et al. 1989). We determined net
seasonal soil-water extraction by calculating the
difference in ASW between sampling dates.
The field capacity for a given soil depth was
assumed to be the highest level of soil water
recorded during the study period. Measurements
used were those taken soon after a precipitation
event or in early spring after snowmelt. Fieldcapacity estimates for the soil profile averaged3S%
(percent volumetric soil water) for Paddockwood,

27% for Floral,3l% for Hagen, 30% for Clair,

24% for Outlook and 38% for Perdue. These

permanent record of the root system. The pictures

were taken at night to improve the contrast between

values are close to previous field-capacity estimates

for Saskatchewan soils (de Jong 1967).

the light-coloured roots and the darker soil. The

exposed vertical face of the trench was illuminated
by three 400-W light bulbs. We prepared each piot
for photography by anchoring a 120 cm x 150 cm
frame to the vertical face of each plot. The frame
contained 30 cm x 30 cm grids, which helped us
identify the location of sample areas in the soil
profile. Several photographs were taken of each
frame to ensure a complete record of the rooting
pattern at each depth.
The pictures of the profile wall were projected
onto a screen with 3 cm x 3 cm grids. The distance between the projector and the screen was
adjusted so that the actual size of the profile wall
was achieved. The intersections of the roots with
the 3 cm x 3 cm grid lines were counted, providing a record for each soil depth (0-10, 10-30,
30-50, 50-70, 70-90, 90-110 and ll0-130 cm).
Four 12 cm x 18 cm samples were used to
estimate average root length for each depth. Root

Maximum root depth was established at each

sampling date as the average maximum depth of
roots for several locations along the vertical trench
face within each plot. Soil-water depletion patterns
were also used as indirect measures of effective
crop rooting depth. In this instance, effective
rooting depth was defined as the lowest increment
(averaged over replicates) in which a significant
(P < 0.05) decrease in soil water occurred
between sampling dates. This soii-water-depletion
method has been found to work best under dry conditions (Bohm 1979). ln the present study, this
method could only be used to estimate rooting
depth to the nearest 20 cm (i.e., one soil-depth
lncrement).
Weather stations equipped with microloggers
(Model CR-21; Campbell Scientific, Logan, Utah)
were located at all sites. Precipitation was recorded
with a tipping bucket rain guage (Model RG2501,
Sierra Misco Inc., Berkeley, CA). Soil temperature was measured at all trials in 1985 and 1986
by thermistors placed at 5, 20, 50, 100 and 120-cm

length was determined with a modified lineintersectmethod (Tennant 1975): rootlength : N*

Dl4, where N represents the number of intersects
and D represents the distance between grid lines.
Volumetric soil water was measured at intervals
during the growing season with a neutron probe

soil depths.
Analysis of variance (Statistical Analysis Systems

North Carolina). We gravimetrically determined

soil-water content at 0-10 cm. then multiolied it
by soil bulk density to convert it to volumetric

Institute, Inc. 1985) was used to determine the significance oftreatment differences. Since only one
replicate per site was used for direct root observations, analysis was conducted with iocations as
replicates. The significance of relationships
between soil-water extraction and the presence of

basis. Available soil water in each of the seven soildepth increments was calculated as total volumetric

roots and between effective and actual rooting depth

was tested by correlation and regression analysis.

(Model 3330, Troxler Laboratories, Triangle Park,

soil water minus the lower limit. The lower limit

was established by the pressure plate method at
- 1.5 MPa. A lower limit was not established for
the Paddockwood site. Data from this study were
expressed as available soii water (ASW) (data for
Paddockwood expressed as total volumetric soil
water). Significant differences in ASW between
treatments were assumed to reflect differences in

RESULTS AND DISCUSSION

Root Growth
Distribution of roots in the soil profile was
more uniform at Perdue in 1985 than in the
1986 trials (Table 1). Greater root length
below 50 cm at Perdue may have been related

ENTZ ET

AL.

to greater availability of soil moisture. The

available soil water to 130 cm at anthesis was
14 cm at Perdue, compared with an average
of less than 5 cm in the 1986 trials.

Can. J. Plant Sci. Downloaded from pubs.aic.ca by 50.26.13.38 on 10/24/14

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The rooting patterns of SW and WW

cultivars were monitored throughout the 1986
growing season. Differences in the rooting
patterns of semidwarf and tall cultivars were
not detected (P > 0.05). Non-significant
differences in root densities of tall and semidwarf cultivars have also been reported by
Lupton et al. (1974) and Cholick et al.
(1977). In contrast, the earlier development
of WW resulted in a greater root length in the
soil prohle by the end of May, and this advantage was maintained until WW maturity

(Table 1).
Both WW and SW cultivars had rooted to
a depth of 50 cm by the end of May (date l)
in 1986 (Table 1). Klepper et al. (1973) also
reported that rooting depth between tillering
and stem elongation in wheat ranges from 30
to 50 cm. Similarly, Campbell et al. (1977)
observed that, by the three-leaf stage, spring
wheat on the Canadian orairies had 62 and

23% of its roots in the top 15 and

the

15-30 cm soil layers, respectively.

A second set of root measurements was
taken at the end of June 1986 when the WW
and SW were at Zadoks growth stage (ZGS)

A third set of root measurements was taken

in mid July 1986 when the WW and SW were
atZGS 89 and 65, respectively. Wheat roots
have been reported to reach their maximum
depth at anthesis (Conor 1975; Bauer et al.

1989). However, rooting depth

of

WW

increased between ZGS 65 and 89

in

1986

(Table 1). Rooting patterns can be highly

modified by soil-water availability (Kmoch
et al. l95l; Proffitt et al. 1985; Barraclough
et al. 1989), and it is probable that this lateseason growth of WW roots was due to an
improvement in water availability during the

latter part of the 1986 growing season

(Fig. 1). Although SW cultivars approached
their maximum rooting depth by anthesis, an
increase in SW root length between ZGS 65
and 90 (Table 1) was also associated with
higher levels of ASW during this period.

WW and SW roots had penetrated to depths

of

110 and 130 cm, respectively, by crop

maturity in 1986. However, 59% of thetotal
root length was still located in the top 30 cm
of the soil profile on the last two sampling
dates (Table l). Similar observations were
made by Wilhelm et al. (1982), who found
approximately 60% of dryland wheat roots
located in the top 30 cm of the soil at
flowering, and Campbell et al. (1977), who
found that root density decreased curvilinearly

65 and 35 (Zadoks et al.1974), respectively.

Root length in the surface 30 cm of the soil
more than tripled between the first and second
sampling dates. with little increase in rooting

with depth.

depth (Table 1).

Bauer et al. (1989) found that the rooting
depths of WW grown on the northern Great
Plains typically were at least 1 m by anthesis
(ZGS 65). In the present study, maximum
rooting depths for WW at anthesis were 110
and 70 cm in 1985 and 1986, respectively
(Table l). Greater root penetration at Perdue
in 1985 could be attributed to higher levels
of preanthesis soil moisture (Kmoch et al.

stage

1957; Proffitt er al. 1985). The ASW in

each 20-cm soil-depth increment between
10 and 130 cm at the boot stage (ZGS 45)
ranged from 1.7 to 2.9 cm at Perdue in
1985, compared with less than 1.0 cm in the
1986 trials.

1113

WHEAT ROOT GROWTH AND SOIL-WATER EXTRACTION

The rooting patterns (especially maximum

rooting depth) of SW and WW were similar
at each sampling date despite differences in

of

development (Table

1).

This

similarity suggests that either (1) factors other

than growth habit had a major affect on root
production or (2) SW roots grow faster than

WW roots.
SW plants start the growing season with an

intact seminal root system, whereas the

seminal roots of WW plants are winterkilled in regions with harsh winter climates
(Chen et al. 1983). Seminal roots have been
reported to produce a more uniform root
distribution than nodal roots (Bedford et al.
1987), which may also allow the rooting depth

of SW to increase more quickly than that

of WW under western Canadian orairie
climatic conditions.

1114

CANADIAN JOURNAL OF PLANT SCIENCE

Soil temperature may limit root growth of

both SW and WW, thereby masking differences in root-growth potential. Marquis
spring wheat has been shown to develop best
at soil temperatures of 16'C, even though
germination occurs at much lower tempera-

E
U
F

tures (Wort 1940). Miyasaka and Grunes

(1990) observed that WW plants grown at

=2
@

U
J
@r

8"C soil temperature had slower growth rates

and less root extension than plants grown at

Can. J. Plant Sci. Downloaded from pubs.aic.ca by 50.26.13.38 on 10/24/14

For personal use only.

16'C soil
0

temperature.

The l5"C

soil-

temperature isotherm was located between the

20- and 50-cm soil depth on the first sampling
date ofthe present study (Table 2). The percentage of roots in the upper 30 cm of soil

=20
F
&10
0

cLAtR 1987

was 100 at Clair, JJ at Floral, and 93 at

Outlook. The upper 50 cm of soil had
warmed to approximately l5'C by the second
sampling date (WW anthesis) and 99% of
both WW and SW roots were found in this
layer. Even at Perdue, where soil moisture
was more favorable, 62% of the root length
was found in the upper 50 cm of the soil profile at anthesis, with another 23% inthe next
soil-depth increment (Table 1). Therefore, the
region of root distribution once again coincided with soil temperatures above 15'C

fr

=2
ql

Or

(Table

2). The relationship

between soil

temperature and root distribution after

anthesis was more difficult to determine

because soil temperature was only measured

cLAtR 1988
E

at 50, 100 and 120 cm. However, evidence

from the preanthesis period suggests that soil
temperature could have influenced rooting
depth of all cultivars evaluated. Further
research is required to better establish the
effect of soil temperature on root growth of
crops produced on the Canadian prairies.

E
U

Soil-water Extraction

Net soil-water extraction (i.e., change in level

of soil water

u
6

between sampling

dates

excluding precipitation) between the start of

spring growth and WW harvest ranged from
1 .3 to 5.8 cm, while net soil-water extraction
between spring and harvest of SW ranged

.0
.20
F

&ro
o

Fig. 1. Available soil water to

a depth of 130 cm
for three growing seasons at Clair, Saskatchewan.

ENTZ ET AL.

Table

2. Soil

temperature

('C) by

depth for experiment sites

in

t9.9

15.0

17.5
18.4
18.1

11.9

68

16.4
18.9
18.6

31

20

aA

)..J

34

15.1

18.1
19.4

11 .3

65

15.3

10.6

146
170

29
64

18

20.0
21.0

18.5

39

191

89

65

19.5

t9.6

13.9
14.9
17.3

12.0
14.1

Perdue

1985

170

65

Clair

1986

154

)/

11'7

198

Can. J. Plant Sci. Downloaded from pubs.aic.ca by 50.26.13.38 on 10/24/14

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1986

cm

19.5

DOYZ

Outlook

cm

20 cm

Year

986

1985 and 1986

5cm

Location

in

Saskatchewan

Soil temperature ("C)

ZGSY

Floral

1115

WHEAT ROOT GROWTH AND SOIL-WATER EXTRACTION

148

l7l

65
84

21
55

19.1

50

100

120 cm

8.2
6.5
0.5

t4.3

i4.8

t2.0

ln

8.7

t2.9

"Day of year.

vZadoks etal. (1974) growth stage of winter wheat (WW) and spring wheat (SW) cultivars.

from -0.1 to 7.5 cm" The one negative value

was attributed to recharge of soil water between
spring and SW harvest in one trial. In the frve
ffials where all four cultivars were included, net

soil-water exffaction from spring to WW harvest averaged 3.'7 cm for Norstar, 2.5 cm for
Norwin, 2.8 cm for Katepwa, and2.9 cm for
l{Y320, while net soil-water exffaction between
spring and SW harvest averaged 2.J cm for

for Norwin, 4.3 cm for

Katepwa and 4.8 cm for HY320. Less soilwater extraction for WW cultivars between
May and SW harvest was attributed to lateseason precipitation that occurred after WW
was mature but while SW was still growing.
Therefore, SW used water while WW did not.
In this study, net soil-water extraction
across all cultivars amounted to approximately
2O% (range 5-35%) of the total ET. This
supports a similar observation for Norstar
WW by Entz and Fowler (1989).
Seasonal patterns of ASW are shown in
Fig. 1. The trends at Clair closely reflected
results at the other sites within years, and
therefore only data from Clair are shown.
Over all trials, maximum soil-water extraction (ASW in spring minus lowest level of
ASW recorded excluding precipitation use)
for the seven soil increments (0-130 cm)
averaged L.2, L.4, 1.0, 0.6, 0.3, 0.2 and
0.2 cm, respectively. Therefore, as reported
Norstar, 2.0 cm

by Doorenbos and Kassam (1979) and Proffitt

et al. (1985), most water extraction for wheat

occurs in the upper 70 cm ofthe soil profile.

Net seasonal soil-water extraction (ASW in

spring minus ASW at harvest excluding

precipitation use) averaged 0.2,0.1 ,0.6, 0.5,
0.3, 0.2 and 0.2 cm, respectively. Slightly
higher soil-water extraction below 70 cm in
1986 than in 1987 and 1988 (Fig. 1) may be
attributed to drier soil conditions near the
surface. Schurmann (1959) and Proffitt et al.
(1985) found that when surface soil-water
supply was low, roots extracted water from
deeper in the soil profile.
A second calculation of water use included
both soil-water extraction and growing season
precipitation. We estimated infrltration of
growing season precipitation by calculating the
amount of water required to bring successive
soils depths to field capacity. Although this
approach is inadequate for determining the
exact distribution of precipitation in the pro-

file, we considered it

adequate

to determine

maximum potential water infiltration. Results

of the analysis indicated that precipitation water

rarely infiltrated below 50 cm into

the

50-70 cm soil depth. It was found that over all

sites and cultivars, an average 95 % of total soil-

water extraction during the growing season

occurred in the top 70 cm of soil. Similar
observations for dryland wheat have been
reported by others (Brown 1971; Doorenbos
and Kassam 1979; Proffitt et al. 1985).
Early in the growing season, WW cultivars
consistently had a lower level of ASW at

1116

CANADIAN JOURNAL OF PLANT SCIENCE

Table 3. Available soil water (cm per soil depth increment) for Norstar and Norwin winter wheat and Katepwa
and HY320 spring wheat grown in Saskatchewan, 1986 to 1988. (Note: Only soil depths where significant differences
between cultivars occurred are shown)

Location Cultivar
Clair

Can. J. Plant Sci. Downloaded from pubs.aic.ca by 50.26.13.38 on 10/24/14

For personal use only.

1986

Date of Year and Soil Depth Increment

DOY

1422

0-10
Norstar 2.2bv
Norwin 2.2b
Katepwa 3.0a
HY320 3.0a

154

10-30

0.7b
0.7b
l.2a
1.2a

178

0-10 10_30
0.6b 0.3b
0.6b 0.3b
1.0a 0.6a
l.la 0.8a

DOY

198

30_50 0_10 10_30

0.4b t.8b 0.8b

0.4b l.i b t.\ab
O.jab 2.0a l.la
0.9a 2.0a 1.2a

221

30_50

0.2c
O.4b

0.6ab
0.8a
231

10-30 30-50 50-70 70-90 90-110 110-130 30-50 70-90 90_110

110_130

Norstar l.3a 0.6b O.1b 0.3ab 0.7a 0.7a 0.4b 0.2b 0.6a
Norwin l.3a 1.0a 0.3a 0.5a 0.8a 0.ia 0.ia 0.3a 0.8a
Katepwa 0.1b 0.3c O.0b O.lb 0.0c 0.0b O.lbc 0.0b O.Ob
HY320 0.2b 0.3c 0.0b O.lb 0.2b 0.1b 0.0c O.tb 0.0b

133
144
0-10 10-30 30-50 0-10
Norstar l.4b O.4b 0.6b 0.1b
Norwin L4b 0.4b 0.'7b 0.7b
Katepwa l.9a
0.9a \.la 0.9a
HY320 l.6ab 0.8a 0.9ab 0.9a
DOY 153
watrous
1986
0-10 10_30
Norstar .2b l.Ob
Norwin |.2b 0.8&
Katepwa 1.7 a 1.6a
HY320 l.8a |.4tt
DOY 170
212
FIoral
DOY

Outlook
1986

30-50 0-10 10-30

1986

Norstar
Norwin
Katepwa
HY320
Clair
1987

Hagen
1987

0.7b
0.6b
0.9a
0.8ab

2.2a
2.0a
1.1b
0.9b

l.ia
l.5a
l.Ob
0.8b

132
160
0-10 0-10 10-30
Norstar 1.8b 0.1b 2.9b
Norwin 1.3c 0.'lb 2.lc
Katepwa 2.0ab 1.2a 3.3b
HY320 2.1a l.la 3.4ab
DOY 125 183 209
0- l0
30_50 30-50
Norstar 0.4b |.3bc I .5bc
Norwin 0.5b 1.9a 2.lab
Katepwa 0.6a 1.2c 1.2c
HY320 0.6a 1.2c l.2c
DOY

30-50

l.la
1.2a

0.4b
0.4b

t67

10-30 30_50 50_70

0.2c
0.1c
0.1ab
0.4b

0.3b 0.0c
0.4b 0.2bc
0.9a 0.4ab
0.6ab 0.6a

211

50-70
0.0c
0.0c
0.3a
0.6a

O.8a

O.ia
O.Ob

0.0b

ENTZ ET AL.

Table
Location Cultivar
Clair
1988

3.

(Continued)

Date of Year and Soil Depth Increment

DOY 150

0-10
Norstar 0.6b
Katepwa l.2a

Paddockwood'
1987

Norstar

Can. J. Plant Sci. Downloaded from pubs.aic.ca by 50.26.13.38 on 10/24/14

For personal use only.

ll17

WHEAT ROOT GROWTH AND SOIL-WATER EXTRACTION

Norwin
Katepwa

HY320

10-30
2.4b
3.4a

DOY 204

30-50
2.9c
4.6a
3.9b
3 .6b

218

30-50

.8b
5.4a
4.0b
4.3b
3

50-70
3.0b
3.9a

3.Ib
3.8a

"Day of year.
vValues within a soil depth increment followed by different letters are significantly different (P

<

0.05).

xTotal volumetric soil water.

depths

of 0-10, 10-30 and

30-50 cm

(Table 3). Greater early season soil water

extraction for WW was likely due to earlier
aerial dry matter production (Entz and Fowler
1991), and hence a greater transpirational
surface area. In one of the eight trials (Outlook, 1986), greater soil water extraction from
the 50 to 70 cm depth by the WW cultivars
carried through until harvest (Table 3). Other
differences between WW and SW included
(l) a significantly lower level of ASW for SW
cultivars between 90 and 130 cm at Clair in
1986, indicating the possibility of deeper SW
root activity; and (2) higher levels of ASW
late in the season for WW at Floral and Clair
in 1986, likely due to recharge from lateseason precipitation.
Like Lupton et al. (1974) and Cholick et al.
(1977), we found that there were few significant differences in soil-water-extraction
patterns of the tall and semidwarf cultivars
compared in our study. Exceptions were in
1986 at Paddockwood on 14 August, where
both Norwin and HY320 had higher ASW at

depth of 50-70 cm than the two tall

cultivars, and at Clair, where Norwin had a
significantly higher late-season ASW than
Norstar for several soil increments Oable 3).
Higher ASW, and hence lower soil-water

extraction by Norwin, may be a reason for

the lower drought tolerance of Norwin compared with Norstar (Entz and Fowler 1990).

However, an exception to this generalization

occurred in early June at Clair in 1987, where
Norwin had a lower level of ASW at a depth
of 30-50 cm, indicating greater soil-water
extraction by Norwin than Norstar in at least
one instance.
Pierce and Rice (1988) suggested that crop
WUE be separated into two components:

(1) soil-water-recovery efficiency and

(2) physiological efficiency. The similarity in

root growth (Table 1) and soil-water extraction (Table 3) between WW and SW in these
trials indicates that higher WUE for WW
(Entz and Fowler 1991) cannot be attributed
to differences in water-recovery efficiency.
The evidence suggests that higher WUE for
WW in the present study was due to a greater
physiological efficiency. For example, WW
used available water earlier in the growing
season than SW (Table 3), at a time when air
temperatures were lower (Entz and Fowler
1991). Previous workers have established that

dry-matter (de Wit 1958) and kernel (French

and Schultz 7984: Entz and Fowler 1989)
production per unit of ET for wheat is greater
at lower air temperatures.

Relationship between Root Growth and

Soil-water Extraction
We compared maximum rooting-depth estimates from the profile-wall and soil-waterextraction methods for Norstar WW using

1l 18

CANADIAN JOURNAL OF PLANT SCIENCE

soil-water-extraction method than with the

profile-wall technique. Greater sensitivity with
the water-extraction method in this case may
have been due to greater replication. The soilwater-extraction technique was also easier to
use and less labour intensive than the orofilewall method. Therefore, for estimates of root
activify (i.e., soil-water extraction) and maximum rooting depth (Table 4), soil-water-

from the four sites where both rootlensth

and soil-water-extraction measurements wJre
data

taken (Table 4). When considered over all

development stages, soil-water-extraction estimates ofrooting depths accounted for 87 % of
the variation in actual rooting depths estab-

Can. J. Plant Sci. Downloaded from pubs.aic.ca by 50.26.13.38 on 10/24/14

For personal use only.

lished from the profile-wall measurements

(Table 4). The strongest relationship between
profile-wall and soil-water-depletion measures
of rooting depth occurred at tillering.
Since it was not possible to estimate the
exact distribution of precipitation in the soil
profile given the available data, we could not
estimate the relationship between root length
and soil-water extraction. However, 95% of
soil-water extraction and 82% of root srowth
occurred in the top 70 cm of the soil profile.
These observations point out that a relationship between root length and soil-water

extraction did not exist and that the vast

majority of root activity (i.e., water extrac-

extraction pattern as measured with a neutron

probe would be the method of choice. This

approach would be especially advantageous
when a large number of treatments are screened
(e.g., cultivars in a breeding program).

The advantage of using the profile-wall

method was that it provided an actual measure
of root distribution in the soil. This was particularly important in the present study
because the root systems of field-grown WW
had not been described for western Canadian

environments.

tion) was confined to the upper portion of the

rooting zone.

SUMMARY AND CONCLUSIONS

In the present study, differences between

cultivars were detected more easily with the

Table

4.

Differences in the rooting patterns of semidwarf and tall cultivars were not detected.

Comparison of direct and indirect estimates of rooting depth for Norstar winter wheat at three development
stages and four locations in Saskatchewan

Direct

Indirect

measuremeil
of maximum
root depth}
(cml

determination of
rooting
depthx

Location

Year

ZGS'

Perdue

1985

31

5'7

60

65

99

82

105

100
110

29

t)

49
52
99

50
70
90

35

65
84

49
95

30
70

31

+-)

of

56

92

9l

Outlook

1986

64

CIair

FIoral

1986

l 986

Direct measurement of root depth (cm)

0.29 + 0.90 (indirect measurement) 12

(cm)

110

50
10
110

0.81

zZadok et al. (1974) growth

stage.
)Rooting depth measured on exposed profile wall (one measurement per replicate).
xEffective rooting depth : lowest
increment in which a significant (P < 0.05) amount of soil water was used
between sampling dates (mean of three to four replicates).

ENTZ ET AL.

Earlier development of WW resulted in

greater early-spring root length for WW than

for SW. This root-length advantage was main-

tained until WW maturity. However, the

rooting depth and pattern within the soil profile were similar for SW and WW at each
sampling date, despite differences in the
stages of development, suggesting that the
zone of rooting was determined more by

Can. J. Plant Sci. Downloaded from pubs.aic.ca by 50.26.13.38 on 10/24/14

For personal use only.

environmental conditions than by genotype or

growth habit. Total root length and rooting

depth increased with level of ASW, and
preanthesis root distribution coincided with
soil temperatures above 15'C. Following
higher early-season soil-water extraction
by WW, only subtle differences in soilwater-extraction patterns between WW and
SW cultivars were observed. As reported
by other researchers (Lupton et al. 1974:.
Cholick et aI. 1977), only small differences

in soil-water extraction were observed

between tall and semidwarf cultivars. Therefore, higher water-use efficiency and grain
yield for WW in these trials (Entz and Fowler
1991) did not appear to be due to differences

in

depth of soil-water extraction between

WW and SW wheat cultivars. However, they
may have been due to differences in time
of water extraction: that is, WW extracted

in the season when growth

conditions favour higher water-use effi ciency.
water earlier

While trends in rooting depth and soilwater extraction were similar, soil-water
extraction was easier to measure, allowing
for more intensive sampling and rigorous
hypothesis testing.

ACKNOWLEDGMENTS
The authors gratefully acknowledge the technical
assistance of K. G. Greer. J. Brydon is aiso gratefully acknowledged for assistance in data analysis.
Financial support for this project was provided by
a grant from the Canada-Saskatchewan Economic
Regional Development Agreement (ERDA).

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