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African Journal of Range & Forage Science

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The effect of defoliation during wilting on the

production of Themeda triandra and Eragrostis
lehmanniana in semi‐arid grassland
a
H.A. Snyman
a
Department of Pasture Science , University of the Orange Free State , P.O. Box
339, Bloemfontein, 9300, Republic of South Africa
Published online: 29 Oct 2010.

To cite this article: H.A. Snyman (1993) The effect of defoliation during wilting on the production of Themeda
triandra and Eragrostis lehmanniana in semi‐arid grassland, African Journal of Range & Forage Science, 10:3,
113-117, DOI: 10.1080/10220119.1993.9638336

To link to this article: http://dx.doi.org/10.1080/10220119.1993.9638336

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 Afr J Range For Sei 1993 10(3)
The effect of defoliation during wilting on the production of Themeda triandra and
Eragrostis lehmanniana in semi-arid grassland
H.A. Snyman
Department of Pasture Science, University of the Orange Free State, P.O. Box 339,
Bloemfontein, 9300 Republic of South Africa
Received 1 August 1992; Accepted 12 July 1993
Abstract
The effect of defoliation during wilting on the production of Themeda
triandra and Eragrostis lehmanniana was studied over four years
(1977 to 1980). Defoliation whilst the plants were at temporary
wilting point in the early growing season (before 1 January) decreased
production from both species in the subsequent season (P 0.01). De-
Downloaded by [North Carolina State University] at 17:16 09 March 2015
foliation over the last half of the growing season, during wilting, had
little effect on growth in the following season (P 0.05). The effect
of a defoliation treatment on production was not necessarily neutra-
lised after one season's rest. Themeda triandra produced more
(P<0.01) than E. lehmanniana for each defoliation. The seasonal
soil-water extraction pattern in the soil profile was also determined.
As the season progressed, plants showed visible signs of wilting at
lower soil-water content than earlier in the season. Most water ex-
traction over the season takes place to a soil depth of 600 mm.
Additional keywords: Drought, soil-water content, temporary wilting
point, Themeda-Cymbopogon veld
Introduction
The fact that the greater part of southern Africa lies in a zone
of arid climate places serious limitations on the agricultural
potential of the region. In South Africa, large parts of the veld
are subjected to moisture stress for long periods due to erratic
and unreliable rainfall. In the extensive grazing areas, with a
mean annual rainfall of 500 mm or less, (about 65% of the
area (Schulze 1979)), water availability is the environmental
factor most limiting to production (Snyman 1989). Drought is
a normal phenomenon in the region (Snyman & Van Rensburg
1990).
By understanding plant responses to drought and to defolia-
tion, it should be possible to cope pro-actively with drought.
Many of the negative impacts of drought on natural resources
are man-induced (Snyman & Fouche 1991). Although an
adequate knowledge of growth patterns, responses to defolia-
tion and to moisture stress in perennial grasses is a prerequisite
for the optimal use of veld for animal production, knowledge
of this subject is limited. In the central grassveld region,
Themeda triandra is the dominant grass species in veld in good
condition, with Eragrostis lehmanniana dominating veld in
moderate condition (Snyman & Fouché 1993). The aim of this
study was to investigate the effects of defoliation during
wilting on the production of Themeda triandra and Eragrostis
lehmanniana in semi-arid grassveld.
113
Procedure
Study area
The research was conducted on a c. 0.2 ha plot at
Sydenham.the experimental farm of the University of the
Orange Free State, five kilometres west of Bloemfontein
(29°06'S, 26°57'E; 1 350 m a.s.l.) which is situated in a
semi-arid region of summer rainfall (annual mean 560 mm,
55% of which falls during the period January to April). The
mean maximum daily temperatures c. 30°C and 17°C in Jan-
uary and July, respectively, but extremes of 41°C in January
and 28°C in July have been recorded. On average, frost occurs
on 119 days each year.
Grassveld typical of the dry Themeda-Cymbopogon veld
(Acocks 1988) was chosen for this study. The soil was that of
the Shorrock series (Hutton form) (MacVicar et al. 1977). Ac-
cording to the new soil classification system (Soil
Classification Working Group 1991) the fine sandy loam soil
is that of the Bloemdal form (Roodeplaat family — 3200). The
three distinct horizons (A: 0—200 mm, B2: 200—600 mm and
IIB2: 600—800 mm) contained, respectively, 10.6, 19.0 and
38.8% clay, and the respective bulk densities were 1 484,
1 563 and 1 758 kg nT3.
Treatments and technique
Twenty-five randomly selected tufts of each of Themeda
triandra, an important decreaser species (Foran et al. 1978),
and Eragrostis lehmanniana, an Increaser Ha species (Fourie
& Visagie 1985) were marked with numbered steel pegs. After
marking, the entire plot was cut to a height of 50 mm at the
end of July 1977. The marked tufts were again cut to a height
of 50 mm during the following growing season (1977/1978),
as soon as they showed visible signs of wilting. A plant was
defined as being wilted when it showed a loss of turgidity
between llhOO and 16h00 (Snyman et al. 1987). In this way,
the temporary and not the permanent wilting point was
measured. After the plants had wilted and had been defoliated,
a new set of plants was identified and marked, and these plants
were defoliated to 50 mm when they showed signs of tem-
porary wilting. This process was repeated for each wilting
throughout the season for each of two seasons viz. 1977/1978
and 1978/1979. A new set of plants was used for each season.
All the marked plants of each wilting over the season, as
well as the rest of the plot, were defoliated to 50 mm at the
end of the growing season, after the plants had become
dormant. The effect of the defoliation (during wilting)
treatments, on plant production during the following growing
season, was determined by allowing the treatment plants a full
 114
growing season's rest in the season subsequent to treatment.
The marked plants were then defoliated to 50 mm once they
had become dormant and their yields (measured in grams)
determined.
After each defoliation the basal area of each tuft was traced
on transparencies, cut out, and sprayed with paint. The areas
were then determined using a leaf area meter. Yield was
2
expressed as mass per unit tuft basal area (g m~ ).
Moisture stress was characterised on the basis of the water
extraction patterns occurring at different soil depths over the
experimental period. Soil-water content was determined
(gravimetrically) weekly for each horizon at depths of 0-200,
200-600 and 600-800 mm. Rainfall and class A pan
evaporation were determined over the experimental period
using standard meterological instruments.
Results
Soil-water content
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During the 1977/1978 growing season, the two grass species
simultaneously showed visible signs of wilting on 5 October
1977, 15 November 1977, and 8 February 1978 (Figure 1). It
was very difficult to identify the wilting order between these
two species. The wilting lasted for periods of between one and
two weeks, but never longer than two weeks.
During the 1978/1979 growing season the two grass species
simultaneously showed visible signs of wilting on 3 November
Depth (mm)
0-800 r—200-800
-600-800
1 Temporary wilting point
H B 2 -Horizon
Ū Rainfall
80-
60-
40-
20-
a
O2-O9-T7 05-10-77 15-11-77 05-02-78 24-03-78
Weeks
Figure 1 Mean weekly soil-water content (mm) and permanent
wilting point (PWP-bars represent the width of the soil-water
content bands for each horizon below which PWP will be
reached), as well as the total weekly rainfall (mm) and class A
pan evaporation (Eo) for the 1977/1978 growing season.
Afr J Range For Sei 1993 10(3)
1978, 30 December 1978, 5 February 1979, and 9 March 1979
(Figure 2). The wilting again lasted for one or two weeks.
Plants marked subsequent to the final wilting period of each
season were defoliated at the end of the growing season (when
dormant) to provide an estimate of total seasonal production
(as control plants).
The total rainfall of 562 mm which occurred during the
1977/1978 season (July to June) was almost the same as the
560 mm long-term average for this area. Only 306 mm
rainfall was recorded during the 1978/1979 season, which is
45% lower than the long-term average for the area. The
466 mm of rain recorded during the 1979/1980 season is only
17% less than the long-term average. The exceptionally high
soil-water content which was present at the start of the
1978/1979 growing season (Figure 2) can be ascribed to the
surplus water carried over from the previous season. During
March and April 1978, 264 mm rain fell, which is 90% more
than the long-term average for these two months.
Most seasonal water withdrawal occurred to a depth of
600 mm (A- and B2-horizons) (Figures 1 & 2). The
B2-horizon (200—600 mm) maintained a high water content
throughout the whole growing season. No great variation in
water content occurred at a depth greater than 600 mm during
the 1977/1978 season. The large variation in soil-water content
which occurred at a depth of more than 600 mm during the
Depth (mm)
0-80O 200-800 .600-800
I Temporary wilting point
fM A- Horizon [O B2-Horizon GnB 2 -Horizon
pyyp PWP PWP
_ A-Horizon \¿a |40
'-UB2-Horizon
0-
0!-O9-78 03-IÍ-78 3O-I2-7B 05-02-79 30-03-79
09-03-79
Weeks
Figure 2 Mean weekly soil-water content (mm) and permanent
wilting point (PWP-bars represent the width of the soil-water
content bands for each horizon below which PWP will be
reached), as well as the total weekly rainfall (mm) and class A
pan evaporation (Eo) for the 1978/1979 growing season.
 Afir J Range For Sei 1993 10(3) 115

plants were in a vegetative growth phase, while Eragrostis

R N Control plants lehmanniana plants were in a reproductive growth phase.
Defoliation at the second wilting took place while all the
• U p f 0 W i l t i n 9 l 1977/78

1
1978/79 grasses were flowering, and defoliation at the final wilting
g j After wiltingj Season LSD (O.OI) when the plants were already seeding.
g 1978/79 Season Í356 A similar pattern with respect to the vegetative and

—į
3OOOn reproductive phases was observed for both seasons up to the

1
a.
first wilting (Figures 3 & 4). During the 1978/1979 season,
however, the subsequent wiltings occurred during the
reproductive development of both species.

i
2000-
Themeda triandra plants that had been defoliated on 5
October 1977 (Figure 3a) and 3 November 1978 (Figure 4a)
had a total above-ground phytomass for the whole season that
¿? loooH Y/£=\ —
= was lower (f<0.01) than that of the other defoliation treat-

!
c
o ments. The fact that during these first defoliations the plants
5 -10 -77

K
8-02-78

were still vegetative with a low leaf area, and therefore

O
in
depended on the withdrawal of growth reserves for a large part
of the season, could account for this lower production.
Downloaded by [North Carolina State University] at 17:16 09 March 2015

i
Themeda triandra plants which were defoliated (during wilting)
2500-
b. later in the season had a total seasonal production that differed
301 220 marginally (P>0.05) from the production obtained from
2000- non-defoliated plants at the end of the season. Eragrostis
lehmanniana, which was in a reproductive phase when
1500- defoliated during the first two wiltings of each season (Figures
I
>
o
.a
1000-
|\\j Control plants
500- [~~l Up to wilting"!
1 9 7 8 / 7 9 Season
After wilting I
77/78 77/78 77/78 77/78 1979/80 Season 1978/79 1979/80
78/79 78/79 78/79 78/79 LSD(O.OI) LSD10.0!)
4000-, a.
Seasons J4II 1161
Figure 3 Mean above-ground phytomass production (g m ) of
2
E 3000- V,
Themeda triandra (a) and Eragrostis lehmanniana (b) obtained
for different wilting periods. This included the production up to
each wilting period when the plants were defoliated and the rest ~ 2000-
of the season (1977/1978), total seasonal production for the 3
•a
subsequent season (1978/1979), and seasonal production of o
1000-
control plants.
a
1978/1979 season, can be ascribed to the poor distribution of o
rainfall during the season. The mean weekly water content of
the A-horizon (0—200 mm) was below permanent wilting point •a
c 2000n J266 Į 375
3
(PWP) for the most part of both the 1977/1978 and 1978/1979
seasons. For the full duration of the 1977/1978 growing
season, the mean weekly water content of the B2-horizon
Į
o 1000-
(200-600 mm) was well above PWP, and that of the IIB2-
horizon (600-800 mm) well below PWP.
Water continued to be withdrawn from the soil profile after
the plants showed visible signs of wilting (Figures 1 & 2).
After the plants had shown signs of wilting for the first time 78/79 78/79 78/79 78/79 78/79
79/80 79/80 79/80 79/80 79/80
during the 1978/1979 season (on 3 November), another 38 mm Seasons
was withdrawn to a depth of 800 mm before it was supple-
mented by rainfall. During this period an exceptionally high Figure 4 Mean above-ground phytomass production (g m 2) of
evaporative demand also occurred. Themeda triandra (a) and Eragrostis lehmanniana (b) obtained
for various wilting periods. This included the production up to
each wilting period when the plants were defoliated and the rest
Above-ground phytomass production of the season (1978/1979), total seasonal production for the
When defoliated at the first wilting (5 October) of the subsequent season (1979/1980), and seasonal production of
1977/1978 growing season (Figure 3) all Themeda triandra control plants.
 116
3b & 4b), had a lower (P<0.01) total seasonal production than
that of the defoliation treatments which occurred later in the
season.
The T. triandra plants that had been defoliated during the first
wiltings, (5 October 1977 and 3 November 1978), produced
40% and 26% less (P<0.01), respectively, during the subse-
quent season than did the control plants. In general, no
differences (P>0.05) in production occurred in the subsequent
season whether T. triandra or E. lehmanniana had been
defoliated later in the season during wilting or not. The
defoliation during the second wilting period on 15 November
1977 (Figure 3b) and 30 December 1978 (Figure 4b) had the
most detrimental effect on the total seasonal production of E.
lehmanniana. These defoliation treatments produced 45% and
48% less, respectively, in the subsequent season than did the
plants that had not been defoliated during wilting in the
previous season.
On average, defoliation of plants during a wilting period in
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the early growing season (before 1 January), decreased
(P<0.01) the production of T. triandra and E. lehmanniana in
the subsequent season by 30% and 38%, respectively, relative
to the control plants. Production of plants that had been
defoliated during wilting in the early growing season (before
1 January), decreased in the subsequent season by 25% and
24% (F<0.01) for T. triandra and E. lehmanniana, respectively,
when compared with plants that had been defoliated at wilting
in the last half of the growing season. Generally, plants which
were defoliated during wilting produced more in total over that
season than did the same plants when defoliated only during
the subsequent season when they were dormant (Figures 3 &
4).
Discussion & conclusion
During this study, it was clear that after the plants had shown
visible signs of wilting they continued to extract water from the
soil, thus confirming that the wilting point (temporary wilting)
of a plant is not so much a function of soil-water availability
but rather a function of the plant itself. It was also clear that
when the plants had grown under a low atmospheric evapo-
rative demand (autumn), the soil dries out to a lower soil-water
content than when the demand is high. The reason for this is
that more time is allowed for water to move through the soil
to the roots. Wilting at a lower soil-water content as the
season progressed can be ascribed to an increase in root mass
during the season. When the atmospheric demand is high
(mid-summer), the grasses wilt temporarily in spite of the fact
that the water content of the soil may still be sufficient for
normal growth. The same trend was found by Venter (1988)
with T. triandra and E. lehmanniana in the central grassveld.
Opperman's (1975) results, (also for the central grassveld)
indicated that T. triandra utilises water more effectively at low
soil water levels than at high levels. This applies not only to
periods between seasons, but also within a season. The same
trend was found by Snyman (1989) with T. triandra and E.
lehmanniana in the same area. According to Du Pisani (1991)
any drought classification based only on rainfall, should be
handled very circumspectly, as little similarity exists between
such a classification and one based on soil-water content.
Afr J Range For Sei 1993 10(3)
Because most water withdrawal during the season only
occurred to a depth of 600 mm (A- and B2-horizons) during
this study, it can be concluded that most grass roots can be
found up to this depth. In Themeda-dom'mated grassveld,
76% of roots sampled to a depth of 800 mm were found in the
top 200 mm of soil (Moore 1990). Tainton (1981) also main-
tained that c. 85% of the root mass of grasses can be found in
the first 150 mm of soil, but pointed out that the efficiency of
the deeper roots should not be underestimated. Studies in the
south-eastern Transvaal also indicated that the deep roots of
grasses, although relatively sparse, appear to play a significant
role in water uptake (Barnes et al. 1991). In the present study
it is clear that water was being extracted to a depth greater
than 200 mm.
It was also significant that the B2-horizon (200—600 mm)
maintained a relatively high water content throughout the
season. Snyman (1985) found, on the same soil form, that this
horizon supplements the water content of the A-horizon. No
great variation in soil-water content occurred in the
IIB2-horizon (600—800 mm) during the season. This agrees
with the findings of Snyman (1988) and Snyman & Fouché
(1991) that percolation of water to a depth of 800 mm occurs
only after exceptionally high rains in these semi-arid
grasslands.
During this study, wilting periods were more or less equal
in duration (< 2 weeks), and thus the general conclusion can be
drawn that defoliation during an early-season drought will lead
to significantly lower production, not only during the remainder
of the growing season, but also for the following season. The
two species studied are not as sensitive to drought during the
last half of the growing season as they are to dry periods
during the first half of the growing season. Pre- and
mid-summer droughts are a normal phenomenon in the
semi-arid areas. In the False Thornveld of the eastern Cape,
T. triandra tolerates defoliation during the dormant season, and
is adapted to no more than modest herbivory (Danckwerts &
Stuart-Hill 1987). According to Danckwerts & Nel (1989) this
species is most productive under a regime of long periods
(within a season) of undisturbed growth, and is particularly
sensitive to defoliation in spring (Danckwerts 1987). In the
eastern Cape, moisture stress reduced the rate of leaf
emergence and leaf longevity on tillers of T. triandra
(Danckwerts & Aucamp 1985).
Results from the current study indicate that the carry-over
effects of defoliation frequencies from one season to another
may have the same, or a greater, effect on plant production
than the treatment within a specific season. Recommendations
based on the results of only one season's research should
therefore be handled with circumspection. These data indicate
that the effect of a defoliation treatment on production is not
necessarily neutralised after one season of rest. The
importance of the correct time of utilisation for grazing with
respect to sustained high production has again been
emphasised. The correct utilisation can be determined by
physiological studies, but usually there is not enough time to
wait for basic research, and therefore management
decision-making often rests on observation and empirical
evidence.
Future studies need to monitor the carbohydrate status and
 Afr J Range For Sei 1993 10(3)
minimum period for recovery of grazed plants that have been
subjected to various periods of moisture stress. In view of the
frequency of drought in semi-arid grassveld, prediction of the
response of grasses to moisture stress after defoliation is of the
utmost importance for optimum and continued production in
these areas.
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