Relative digestibility of Agropyron smithii and Bouteloua gracilis grown under six water regimes
by Jeffrey Martin Welker
A thesis submitted in partial fulfillment of the requirements for the degree of MASTER OF SCIENCE
in Range Science
Montana State University
© Copyright by Jeffrey Martin Welker (1982)
Abstract:
Rangeland production needs to be evaluated on a livestock digestibility basis as well as on gross
dry-mater yield. The forage digestibility of two eastern Montana range sites were studied. The first site
was dominanted by western wheatgrass, (Agropvron smithii Rybd) and the second site was dominanted
primarily by blue grama, (Bouteloua gracilis (H.B.K.) Lag. ex Steud). At each site, plots were irrigated
in the preceding fall(20cm), current spring (20cm), and summer (0, 6, 12, 25mm/week). Forage
samples were collected in June, July, August and November in 1977 and June, July, August and
October 1979. Digestibility was compared between water treatments by an in-vitro technique. Results
suggest that while digestibility declines with season in all treatments, for both species, water
availability has little effect on in-vitro digestibility. ■STATEMENT OF PERMISSION TO COPY
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fulfillment
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requirements for an advanced degree at Montana State University., I
agree that the Library shall make it freely available for inspection.
I further agree that permission for extensive copying of this thesis
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Signature _
Date
I^
^VVl. f
T
y
Jy
a
)
RELATIVE DIGESTIBILITY OF
AGROPYRON SMITHII AND BOUTELOUA GRACILIS
GROWM UNDER SIX WATER REGIMES
by
JEFFREY MARTIN WELKER
A thesis submitted in partial fulfillment
• of the requirements for the degree
of
MASTER OF SCIENCE
in
Range Science
Co-Chairman, Graduate Committee
Co-Chairman, Graduate Committee
'TVL
Head, Major Department
^
Graduate Dean
MONTANA STATE UNIVERSITY
Boseman, Montana
June,.1982
iii
ACKNOWLEDGEMENTS
I wish to express my sincere appreciation to the following:
Dr. Tad Weaver for his professional guidance, inspiration, enthusiasm
in plant ecology,
friendship,
and review of the thesis while serving
as a co-chairman of my committee.
Dr. Clayton Marlow for his guidance, friendship, insight into range
science and review of the thesis while serving as a co-chairman of my
committee.
Dr. Kris Havstad for his patience, encouragement, scrupulous review of
the thesis and unique insight into the field of range science.
The many professionals in the Animal and Range Science Department and
the Biology Department, for without their support and encouragement,
this research would have been extremely difficult.
The fellow graduate students with whom I have shared so much time
enjoying and discussing the many aspects of range science.
I would also like to thank my parents,grandparents and other members
of my family for their patience and quiet support during the many
years of my formal education.
Finally, I would like to thank my wife Molly for her devoted patience,
guidance
and
friendship
during
the
duration
of
this
research.
TABLE OF CONTENTS
PAGE
VITA ....................
ACKNOWLEDGEMENTS ................... ".................... .
TABLE OF CONTENTS..................................... ...... .
LIST OF T A B L E S ........ ....... .............. ................
LIST OF F I G U R E S ............. .......................... .
ABSTRACT......
ii
iii
iv
v
vi
vii
INTRODUCTION ...........
I
REVIEW OF LITERATURE................. ........................
4
METHODS .....................
Site Description.......
Water Treatments ............
Forage Sampling..............
Digestibility Trial ........ ............ ............
Statistical Analysis ..............
R E S U L T S ...........
1977
1977
1979
1979
Observations,Aeroovron smithii.................
Observations, Bouteioua gracilis........... ..
Observations,Aeroovron smithii................
Observations, Bouteloua gracilis..... .........
10
10
11
13
15
16
18
18
21
22
23
DISCUSSION AND CONCLUSION ................
26
SUMMARY ......................................................
30
APPENDICES
31
.................. ............... .......
1
Soil Description..................
32
2
Computer Program to Calculate Digestibility and
Data File ..............
33
LITERATURE CITED
41
V
LIST OF TABLES
Table
1
2
3
4
5
Page
Leaf-Stem Ratios of Agroovron smithii and
Bouteloua gracilis Under Six Water Regimes .......
6
Precipitation and Average Temperature Miles City,
Montana 1976-1977 and 1978-1979 .........
14
Forage Sample Collection Dates for Agroovron
smithii and Bouteloua gracilis......
15
In-Vitro Digestibility Trial Results by Laboratory
Trials .............
17
Relative Digestibility of Agroovron smithii
and Boutloua gracilis Under -Six Water Regimes
and Two Years (Mean and Standard Deviation)
19
vi
LIST OF FIGURES Figure
I
2
,,
Page
Phenological Development of Aeroovron
smithii and Bouteloua gracilis ..................
7
Relative Digestibility of Agroovron smithii
and Bouteloua gracilis Under Six Water Regimes
at Four Sample Periods ..........................
20
vii
ABSTRACT
Rangeland production needs to be evaluated on a livestock
digestibility basis as well as on gross dry-mater yield. The forage
digestibility of two eastern Montana range sites were studied. The
first site was dominanted by western wheatgrass, (Agroovron smithii
Rybd) and the second site was dominanted primarily by blue grama,
(Bouteloua gracilis (H.B.K.) Lag. ex Steud). At each site, plots were
irrigated in the preceding fall(20cm), current spring (20cm), and
summer (0, 6, 12, 25mm/week). Forage samples were collected in June,
July, August and November in 1977 and June, July, August and October
1979. Digestibility was compared between water treatments by an invitro technique.
Results suggest that while digestibility declines
with season in all treatments, for both species, water availability
has little effect on in-vitro digestibility.
INTRODUCTION
Western rangelands are a major source of forage for livestock
production.
lands
In order that the animals using
the forage on these
obtain profitable weight gains and maintain reproductive
capabilities, the forage must provide essential nutrients in adequate
amounts and the forage must be digestible by the rumen micro-flora.
Therefore,
any management practice which would improve nutrient
content or increase forage digestibility would prove beneficial to the
livestock producer.
Examples of management practices which may be incorporated to
improve total nutrient
inter seeding,
content
reseeding,
of r a n g e l a n d ’s are:
waterspreading,
fertilization (Vallentine 1980, Bokhari 1978).
brush
chisling,
removal
and
Yet a management tool
which would increase forage digestibility has yet to be implemented.
Eck et al (1981) attempted to increase forage digestibility, with
water supplementation.
(Festuca arundinacea
In the Southern Great Plains, tall fescue,
Scherb.) and smooth bromegrass,
(Bromus inermis
Leyss.) were subjected to three water regimes. In two of the regimes
water was applied
over the entire growing season, while in the other
regime supplemental water was applied only during the cool portion of
the growing
season.
Only
total
period
amounts
of
water
were
2
reported.
This
research
showed
that
there was
no
differences in digestibility across water treatments.
appears
that climatic parameters,
temperatures,
relative humidity,
such as,
significant
It therefore
photo-period,
rainfall pattern;
daily
or species
chracteristics are limiting the ability of these species to respond
with increased digestibility under supplemental water in the Southern
Great Plains.
With the precipitation pattern, daily temperatures, relative
humidity and wind patterns varying from the Southern Great Plains to
the Northern Great Plains (Humprey,
1964).
And with species to
species variation in digestibility (Cogswell and Kamstra 1976 ), the
possibility exist that different species,
in other locations,
respond differently in terms of digestibility
With an
might
to supplemental water.
irrigation project in eastern Montana directed at assessing
the response of two grassland ecosystems to different water regimes,
the opportunity to investigate the relationship between water regimes
and digestibility presented itself.
There were two objectives of this
study.
I)
Determine whether summer irrigation of 0, 6, 12, or 25mm/wk
would
significantly alter the forage digestibility of western wheatgrass,
(Aeroovron smithii Rybd) and blue grama.(Bouteloua gracilis(H.B.K.)
Lag. ex
Steud) and
3
2)
Determine whether water applied at different seasons (Fall,
Spring and Summer) would significantly alter forage digestibility of
these two species
LITERATURE REVIEW
Aeropyron smithii is an abundant native mid-grass found primarily
in the mixed-grass and short-grass prairies of North America (Kuehler
1964).
This species initiates growth in the cool part of the .growing
season, April and May, and achieves most of its production before the
hot raid-summer months (Erickson, 1966).
Plants of western wheatgrass
reach maturity with the production of seed during the early to mid­
summer months, depending upon environmental■conditions.
Bouteloua gracilis is
a warm season grass which usually begins
growth in the warmer summer months of June and July, yet growth may
begain as early as April in Montana.
Maximum growth usually occurs in
late July to late August, yet this period may vary from the southern
ecotypes to the northern ecotypes (Launchbaugh and Hackerott, I969;
Pieper
et al, 1971).
Maturation occurs with the formation of seed,
which usually occurs during the later part of the growing period.
(Launchbaugh and
Hackeroff, 1969).
Both Agroovron smithii and Bouteloua gracilis
are known to be
highly digestible early in their vegetative growth stage (Cogswell and
Kamstra,
1976).
Digestibility of both species decreases as the forage
matures with the progression of the growing season.
are
most
i nfluential
in
dictating
digestibility are leaf-stem ratio,
the
The factors which
seasonal
phonological
changes
stage,
in
species
5
characteristics and concentration of metabolic compounds (Cook and
Harris 1968, Kamstra et al 1968, Kamstra 1978, Erickerson et al 1978).
A primary
result of water supplementation
on native vegetation
in the Northern Great Plains is to increase forage production ( Monson
and Queensberry, I958; Branson, I956; I965; Smika et al, 1965; Perry
1978).
One
wonders
whether
digestibility may also be affected?
leaf-stem
ratio
important, to
Weaver and Welker (in prep.) have
shown that.application of large amounts of water increase leaf-stem
ratios in Aaroovron smithii
communities and barely increase leaf-stem
ratios in Bouteloua aracilis communities (Table I). • Yet the degree of
response of the vegetation to water supplementation may be limited by
other factors; nutrient supply, pattern of application and genetic
limitations (Perry 1976).
Another correlate of digestibility,
be
phenology,
influenced by water supplementation.
Dickinson and Dodd (1976),
Wight and Black (1974), Weaver (1975) and Collins
have all d e m o n s t r a t e d
development (Fig.
I)
is not thought to
this relatioship.
and Weaver (1978)
The phenological
of Aeroovron smithii and Bouteloua gracilis at
our sites in the Northern Great Plains, under a water gradient and
seasonal
w a t e r •additions,
were not significantly influenced
by
additions of water (Weaver and Welker in preparation).
Forage digestibility research has demonstrated that at a given
period different species exhibit differences in digestibility (Cook
6
Table I. Leaf-Stem Ratios of Agroovron smithii and
Bouteloua gracilis under Six Water Regimes in 1980.
Agroovron smithii
Date
25 June
15 July
18 August
28 August
3
8
0.3
3
1.0
3.0.
0.16
88
2.5
0.4
1.9
3.6
9.0
24.0
9.4
24.0
2.6
0.5
1.0
2.6
4.2
2.2
1.8
5.5
1.8
3.3
2.3
. 3.8
2.0
2.1
2.0
1.3
2.4
2.1
Water
Regime
Control
a
6mm/wk
a
I2mm/wk
a
25mm/wk
a
Fall Wet
b
Spring Wet
5.0
5.5
b
Bouteloua gracilis
Control
2.0
2.0
a
6mm/wk
2.5
5.7
a
I2mm/wk
4.2
1.6
a
25mm/wk
4.1
7.0
a
Fall Wet
2.2
1.8
b
Spring Wet
4.4
4.6
b
*= 4 leaf hits, O stem hits
**= 12 leaf hits, O stem hits
a) Minimum weekly water additions, natural + artificial. Control
regime received no supplemental water.
b) Fall irrigation (Sept. I to Sept. 15) 20 cm of supplemental water or
the amount of water required to saturate the soil to 75cm; Spring
irrigation (May 15 to June I) 20cm of supplemental water or the amount
of water required to saturate the soil to 75cm. Fall or Spring were
the only periods these plots received supplemental water.
7
Figure I. Phenological Development of Agropyron smithii and
Bouteloua gracilis.
Agropyron sm ithii
DISSEMINATION
AN THESIS
HEAD
BOOT
---- !977
---- 1979
VEGETATIVE
JUNE
OBSERVATION
------JU L Y -----3
15
22
AUGUST
DATES
Bouteioua gracilis
DISSEMINATION
R /D
ST
RIPE
0 .9
A /R
ANTHESIS
- j 2.
5|
H/A
HEAD
B /H
BOOT
V /B
/977
VEGETATIVE
— AUGUST —
7
OBSERVATION DATES
16
24
8
and Harris 1968;
Kamstra, 1973;
variations observed are
generally
phenological stages of each
growing season.
Cogswell and Kamstra11976).
attributed
to the
The
different
species during a given period in the
Therefore species which complete their life cycle by
mid-July should have a lower digestibility than
species which are
initiating vegetative growth at the same period (Kamstra,
1973).
Digestibility has been shown to be closley related to lignin
content.
Dehority et al (1963)
examined the
relationship, between
lignin concentration and forage digestibility, concluding that as the
plant matures the concentration of undigestible lignin increases.
He
also showed that the lignin polymer acts as a coating agent along the
plant cell wall, perhaps protecting the cellulose molecules arid other
compounds from ruminant micro-flora degradation.
Increasing soil
moisture may indeed increase growth (Perry, 1976), yet alteration of
the genetic coding for specific metabolic reactions, such as those
that might influence digestibility, is not feasible over a short
period of water enrichment (Perry,1976).
Eck et al (1981) i m p l e m e n t e d
supplemental water regimes,
a project
incorporating
two
50 and 100cm of additional water applied
over the growing season, on two introduced species of grasses in the
Southern Great Plains.
The results of this research suggest that
supplemental water applied to these two species in the Southern Great
Plains is not a effective management tool for increasing forage
9
digestibility,
or a tool
for prolonging the period of maximum
digestibility.
Another concern associated with irrigated forage and digestibility
is the affect of season of
water additions.
rainfall or irrigation has been
1976).
The effect of timing of
neglected in range research (Perry,
Although the timing factor has been noted as important in
water use efficiency (Wight and Black,1974), nutritive levels of grass
(Rogler and Haves, 1947) and community composition (Albertson and
Toraaneck,
1965); only Eck et al (1981) tested the effect of additional
spring moisture on forage digestibility.
No effect was observed. The
effect of supplemental soil moisture applied in the fall on native or
introduced grass digestibility has not been assessed.
Methods
Site Description:
During the summer of I 977, an irrigation study was initiated by
Dr. T. Weaver of the Biology Department at Montana State University.
The purpose of this study was to examine the effects of additional
moisture,
which may occur due to weather modification,
vegetation of the Northern Great Plains.
on the
The study sites were on the
USDA Range and Livestock Experiment Station at Miles City, Montana.
Two important grassland ecosystems within the Northern Great Plains
were chosen for the
wheatgrass,
project. One study site was dominanted by western
while the other study site
was dominanted by blue grama.
The sites were located on rarely flooded
and Tongue
Rivers.
cattle grazing for
plains of the Yellowstone
The western wheatgrass site
received light
two years prior to the start of the study
The
blue grama study site has been used for the last 10 years as late
summer and early fall grazing for cattle.
The blue grama site
received relatively heavy cattle use because a windmill and watering
tank were nearby.
Both the western wheatgrass and the blue grama
sites were fenced at the start of the irrigation project to eliminate
any domestic animal interference during the duration of the project.
The blue grama site was approximately 150 meters by 50 meters while
the western wheatgrass. study site was
meters.
approximately 150 meters by 300
Soil orders for the western wheatgrass and blue grama sites
11
are Borollic Camborthid and Ustic Torriorthent ,respectively . (Soil
description are found in Appendix I).
Water Treatments. '
The experimental design was a completely randomized block.
At
each site, six water treatments were implemented, each treatment having
two replications per site.
of which a centered,
Each treatment plot was 20 meters square,
14 meter square staked plot was delineated with
nylon line a t t ached to iron stakes.
The w a t e r
treatments
implemented were applied at three seasons: DSummer water, consisting
of four treatments,(a) a control plot which received no additional
water, (b) a plot which was guaranteed 6 mm/week, (c) a plot which was
guaranteed 12 mm/week and (d) a plot which received at least 25
mm/week or whatever amount was needed to keep the soil profile
saturated to 75 cm, this plot will often be, referred to as the "Wet11
water
treatment
plot..
2) Fall
water, this treatment involved
saturating the soil profile to a depth of 75 cm in
mid-September and
thereafter received no additional supplemental water.
3) Soring
water, in this treatment water was applied in 2-5cm intervals in May
and early June, to wet the soil to 75cm and prevent dessication of the
surface horizon (0-25em) during this period.
The summer irrigation plots were guaranteed their respective
amounts by recording naturally occurring rainfall on the sites, and
12
then determining whether the natural rainfall met the specific water
regimes required for the individual treatments.
short of the water regime requirement,
to meet the
supplemental water was applied
water regime requirements.
exceeded the specified water regimes,
If the rainfall fell
If the natural rainfall
this was recorded in the data
log books, with each week a single unit, such that the previous week’s
precipitation had no bearing on the current weeks’ water criteria.
Water applications were made in early moring windless conditions,
using Rain Jet (Rain Jet, Burbank, Calif.) sprinklers (66u openings)
on 50 cm risers.
Four sprinklers were evenly placed in the plots to
obtain even distribution throughout the 14 meter square treatment
plots.
Nine wedge gauges were placed regularly along the edges and
within the treatment plots to record additions of supplemental water.
A mean value of all nine wedge gauges were used to determine the
amount of water applied to a specific treatment plot.
The water
supply for the western wheatgrass site was a irrigation canal filled
with water from the Yellowstone River.
The water supply for the blue
grama site was from the nearby Tongue River.
13
Forage Sampling.
Forage
samples
were
collected for
all years
(1977-1981) at
approximately monthly intervals during June, July, August, October or
November,
from both sites and all treatments.
The samples were used
to determine above ground production of the primary species.
The
forage samples were taken using 10 randomly placed frames (25x50em)
per water regime. Live and dead material of each species was
clipped
to the ground with shears, separated as to grass or forbs, bagged as
such and dried to constant weight at 35 degrees Centigrade.
The
samples were weighed for biomass determination and then ground
through a 40 mesh screen using a Wiley Mill.
then placed in plastic jars,
The ground samples were
sealed and stored in a dark,
cool
basement for further use.
For this digestibility study, forage samples from 1977 and 1979
were used. These years were choosen because they were similar in
climatic conditions, primarily total precipitation on a water year
basis (Table 2).
This study used two subsamples from each replication
of the six water regimes^ Ia & 1b, 2a & 2b. Therefore, four samples
from each water treatment were analyzed for each month, both species
and for the two years. The actual collection dates are shown in Table
3
Table 2. Precipitation (mm) and Average Temperature (Degrees Celsius) Miles City,
Montana 1976-77 and 1978-79.
Precipitation
Oct.
Nov.
Dec.
Jan.
Feb'.
Mar.
Apr.
May
Jun.
Jul.
Aug.
Sept.
Total
Normal
18.0
13.0
12.1
12.4
12.9
8.1
31.2
52.3
84.3
39.4
30.4
30.2
344.3
1976-77
19.1
4.8
6.9
17.2 ■ 2.5
24.6
6.1
62.23 35.0
48.5
57.4
48.5
332.8
1978-79
6.9
55.1
16.0
8.4
28.9
6.6
19.3
34.5
19.5
70.8
17.0
0.8
283.8
Normal
9.3
0.2
-5.5
-9.3
-5.7
-1.0
7.4
13.5
18.2
23.5
22.5
15.5
1976-77
6.7
-1.4
-3.6
-13.5
-0.16
2.3
10.6
16.3
21.7
23.5
18.7
15.1
1978-79
9.2
-5.1
-10.6
-17.6
-13.3
-1.2
5.2
12.0
19.6
23.6
22.2
18.7
Temperature
Ir"
-P'
15
Table 3« Forage Sample Collection Dates for Agroovron smithii and
Bouteloua gracilis in 1977 and 1979.
1977
1979
June
July
Aug.
Nov.
June
July
Agroovron
smithii
23
23
18
10
21
19
20.
7
Bouteloua
gracilis
I
28
20
10
28
30
24
6
Species
Aug. ' Oct
Digestibility Trial.
The two stage in-vitro technique (Tilley and Terry,
used
1963)
was
to determine the relative digestibility of western wheatgrass
and blue grama at the four dates under the six water regimes for 1977
and I 979. Actual in vitro procedure was performed following Moore
and Dunham (1978), with
inclusion of phosphate buffer
as recommended
by Larry White (personal communication with K. Havstad, 1981).
A rumen fistulated black angus heifer was used as the inoculum
source. The animal had been fed a grass hay diet for five days
prior
to the inoculum removal for the digestion trial.
Two digestion trials were run, one for each year.
limitations each trial was split in-half,
Due to lab
such that half of the
samples were cooled for 6 hours the other half cooled for 50 hours.
16
There was no significant difference
trials (Table 4).
(P<.01 ) between these half
Dry matter of each sample was determined to
incorporate into the digestibility calculation (Moore and Dunham,
1978).
Statistical Analysis
Mean
digestibility
values were
calculated and analyzed for
significant differences at the 95% level using an Analysis of Variance
Multi-factor.
Years,
significant F values
species,
months and treatments were tested for
17
Table 4. In-Vitro Digestion Trial Results grouped by Laboratory
Trials.
1977
Laboratory Trial
X
Ia
50.73
SD
SE
8.17
0.83
8.40
0.85
7.04
0.71
5.70
0.58
a
Ib
51.38
a
1979
2a
48.70
a
2b
50.00
a
a, similar sub-script letters indicate no significant difference
between mean values with in each year (PC.01) Student t-test.
Results
1Q77 Observations.
Agroovron smithii.
In 1977 the
digestibility of Agroovron smithii
the control water regime was 57*4# in June.
progressed
diges t i b i l i t y
of
this for&ge
forage under
As the growing season
declined,
with
the
digestibility being 51.0%, 47.3% and 43.0% during July, August and
November,
respectivly (Table 5, Fig. 2).
Water additions of 6, 12, or 25 mm/week during the summer months
did not significantly alter the forage digestibility of Agroovron
smithii
(Table 5).
Digestibility of the watered forage did follow
the same seasonal trend as that of the control forage,
exception,
that
being a slight
but
insignificant
with one
increase
in
digestibility of the 25 mm/week Agroovron smithii treatment between
June and July (Table 5, Graph 2).
Seasonal water addition to Agroovron smithii forage was unique in
1977, such that no water was applied in the fall of I 976, yet water
was applied in the spring of 1977.
Therefore the fall water treatment
plots were a duplicate of the control treated forage, until the fall
of 1977 when the designated fall water plots of Agroovron smithii
received approximately 10 cm of water, or enough to saturate the soil
profile to the 75 cm level.
The response of the Agroovron smithii
19
Table 5
Relative Digestibility (%) of Agropyron smithii
and Bouteloua gracilis Under Six Water Regimes and Two Years
(Mean and Standard Error).
Agropyron smithii
Water
Regime
O mm
week
12 inn
week
week
Fall
Wet
Spring
Wet
Wet
1977
1979
1977
1979
1977
1979
1977
1979
1977
1979
1977
1979
57.3
1.7
54.0
2.1
57.0
0.7
57.0
0.7
56.0
0.4
58.3
1.6
55.8
1.7
58.3
3.8
52.5
5.8
59.8
2.0
55.5
0.8
55.8
2.9
Julyb
51.0
0.9
50.8
2.2
54.3
0.9
50.0
0.6
53.0
1.4
52.8
0.3
52.0
1.2
49.3
0.6
58.3
1.8
49.5
1.3
57.3
0.5
52.3
1.5
Auguetb
47.3
1.3
53.3
1.3
50.8
1.8
49.0
1.7
51.8
0.5
50.3
1.0
50.3
1.8
50.3
3.1
56.8
0.3
46.8
1.5
53.3
3.9
49.8
1.5
Oct./Nov.c
43.0
2.2
49.3
2.0
40.3
1.3
46.8
1.2
38.8
2.2
47.3
1.7
57.0
3.8
46.3
1.3
36.0
4.3
45.3
1.7
41.8
1.3
47.8
1.4
Months
i
June®
Bouteloua gracilis
Water
Regime
0 mm
week
Fall
Wet
12 mm
week
week
Spring
Wet
Wet
Months
1977
1979
1977
1979
1977
1979
1977
1979
1977
1979
1977
1979
June a
60.5
2.3
55.5
2.5
60.5
1.9
55.5
2.7
62.3
1.0
54.0
1.6
62.5
1.2
52.0
0.9
62.3
0.9
53.0
1.8
61.0
1.2
58.0
2.2
July b
56.8
1.3
47.5
1.7
56.5
1.2
51.0
1.2
56.1
1.3
47.0
0.9
56.0
0.9
41.8
3.8
52.8
0.5
50.0
1.0
52.0
2.0
49.3
1.3
August b
45.8
1.0
46.0
2.0
43.8
1.5
49.3
2.7
46.0
1.3
45.8
1.1
46.8
1.8
48.5
1.3
43.8
1.3
45.3
1.9
42.5
2.7
42.8
1.5
Oct./Nov.®
43.5
3.1
46.5
1.5
41.0
2.6
44.0
1.7
38.8
1.1
40.0
0.4
56.0
2.2
45.0
1.7
38.5
4.2
40.8
1.8
38.3
1.1
44.3
1.0
a,b,c superscripts indicate significant differences at the
99% level when years, species and treatments were pooled.
20
Figure 2. Relative Digestibility of Agropyron smith!! (AGSM)
and Bouteloua gracilis (BOGR) Under Six Water Regimes at Four
Sample Periods.
AGSM 1977
jO (C O N T R O L )i 3 m
II g m m / W K
m /WK
AGSM 1979
FI°
LI I g
K M LL W E T i SPW tNO W T T
(C O N T R O L ). 9 m m / W K
m m / WK
Q F*LL WETiSFmio WET
UUL
MONTHS
BOGff 1977
21
forage to the first year of fall watering
was
an increase in
relative digestibility of 7% over that of the previous sampling
period.
Spring watering of Agroovron smithii enhanced relative
digestibility of the forage.
In June the digestibility was 52.5%,
with an insignificant increase in digestibility by July to 58.3%. The
data for Spring Wet
digestibility
of
(Table 5) s h o w s
the
control
forage
that
was
in August
9.4%
less
of
1977
than the
digestibility of spring watered Agroovron smithii. By November,
the
control forage digestibility was greater than the Spring Wet forage
digestibility.
1Q77 Observations. Bouteloua gracilis.
Water additions applied to Bouteloua gracilis forage during the
summer of 1977 (0, 6, 12, or 25 mm/week)
the forage
digestibility.
did not significantly alter
Regardless of summer water treatment,
Bouteloua gracilis forage digestibility was high in June (60-62%), but
declined there after by 0.1 - 0.2%/day regardless of water regime.
Though digestibility
was on a downward trend in late July,
the
digestibility of the Bouteloua gracilis forage was still above 50%
(Table 5).
below
50%,
By late August summer watered Bouteloua gracilis dropped
with values ranging between 42.5% to 46%.
Relative
digestibility continued to decline from late August to early November
regardless of summer water regime.
Seasonal water irrigation applied to Bouteloua gracilis forage in
22
the
spring
of
1977
did
not alter
the forage
digestibility.
Digestibility of the spring watered forage followed the digestibility
trends of the control forage quite closely, falling slightly but
insignificantly below the control forage digestibility in November
(Table 5 and Fig. 2).
The fall watered Bouteloua gracilis forage was the same as the
control treated forage until September of 1977,
at which time water
was applied to the designated treatment plots of Bouteloua gracilis.
As noted at the Agroovron smithii site, relative digestibility of
Bouteloua gracilis increased between August and October in the fall
watered plots in 1977.
probably
This forage digestibility increase was
associated with fall regrowth of both BouteloUa gracilis and
Agroovron smithii after fall watering.
IQ7Q Observations. Agroovron smithii.
In 1979
relative digestibility of Agroovron smithii was less than
60%, regardless of summer water regime in the month of June (Table 5).
Values for the 0, 6, 12 and 25 mm/week water treatments were 54%, 57%,
56.8% and 55.8%, respectively.
A general seasonal decline in forage
digestibility for these treatments occurred from June through the
early part of N o v e m b e r
(Table 5).
By N o v e m b e r
the relative
digestibility of Agroovron smithii, regardless of summer water regime,
declined to between 45 and 50%.
23
Fall
watering of Agroovron smith!! produced a statistically
insignificant increase in digestibility
over that of the control
forage in the month of June.
Values for these treatments in June were
58.3%, and 54%, respectively.
As the 1979 growiiig season progressed,
the fall watered Agroovron smithii digestibility paralleled the
control forage digestibility, through July, August and November (Table
5).
The Fall Wet forage treatment plot received
yearly watering in
September of 1979» but the Agroovron smithii forage digestibility
after this treatment did not increase between August and November as
noted in I977 (Table 5).
Spring watering of Agroovron smithii in 1979 increased forage
digestibility over that of the control digestibility, 59.8% and 54%
respectivley.
After
the month
of June,
the Spring Wet
forage
digestibility followed the digestibility trends of the control forage
until the November sampling period. At this time the spring watered
■forage was insignificantly lower at the end of the season than the
digestibility of the forage under the control water regime (Table 5).
1Q7Q Observations. Bouteloua gracilis.
The digestibility of summer watered Bouteloua gracilis forage was
similar in the month of June with one exception.
For
the Wet water
regime, the relative digestibility was 58%, this was not a significant
difference.
Digestibility of all the summer watered treatments
declined as the growing season progressed (Table 5).
The largest
24
decrease in digestibility on a monthly basis occurred between June
and July regardless
of summer water
regime.
By
November
the
digestibility of summer watered Bouteloua gracilis forage decreased
below 50% digestibility, with the 12 mm/week water regime having the
lowest at 40% (Table 5).
Fall watered Bouteloua gracilis forage digestibility was very
similar to the control forage digestibility in the month of June 1979.
Between June and July the digestibility of the fall watered Bouteloua
gracilis decreased dramatically from 52% to 41%.
Between July and
August, however, digestibilty of fall watered Bouteloua gracilis
increased from 41% to 48.5%.
From August to November the forage
digestibility of the fall watered Bouteloua gracilis decreased to 45%.
The yearly September watering of these treatment plots occurred in
1979, yet as noted with Agroovron smithii. forage digestibility of
Bouteloua gracilis did not increase between August and November, as it
did in 1977.
Spring watered Bouteloua gracilis forage digestibility was
similar to the forage digestibility of the control treated forage in
the month of
June at 53% and 55.5%, respectively.
As the growing
season progressed relative digestibility of spring watered Bouteloua
gracilis forage declined, such that the digestibility in July, August
and November was 50%, 45.3% and 40.8%, respectivley.
By November the
25
digestibility of the spring watered Bouteloua gracilis forage was 6%
less than the digestibility of Bouteloua gracilis forage under the
control water regime, yet this difference was insignificant (Table 5).
The only significant interaction
was that of monthly differences
in digestibility, when years, species and treatments were pooled.
DISCUSSION & CONCLUSION
The principle observations from
this study were:
I) in all
treatments
relative digestibility of Aeroovron smithii and Bouteloua
gracilis
declined from
June
through
November,
2) relative
digestibility of summer watered forage was not increased, 3)
fall or spring water
digestibility, and
additions
significantly
neither
altered forage
4) digestibility decline with season
was not
postponed with water supplementation.
Previous
r e s earch
has
shown
seasonal
changes
in forage
digestibility (Cook and Harris, 1968; Kamstra, 1973; Cogswell and
Kamstra,
1976).
These seasonal fluctuations in digestibility of native
forage have been attributed to numerous factors; leaf-stem ratios,
phenological stage and concentration of metabolic compounds (Cook,
1968; Dehority et al, 1963; Kamstra, 1973).
A significant portion of
these seasonal changes in digestibility have been ascribed to the
lignification of the plant tissue as the maturation process occurs,
with an inverse relationship developing between digestibility and the
lignin content of the forage (Dehority et al, 1963).
•this inverse relationship may
The existence of
make alteration of digestibility
difficult.
Control of forage digestibility must involve both genetic and
environmental parameters.
This study examined the environmental
27
parameter of various water regimes and noted no significant effect on
forage digestibility as well as no effect on seasonal decline in
relative digestibility of Aeroovron smithii and Bouteloua gracilis.
These findings are supported by the work of Eck et al, (1981) who made
similiar observations on smooth bromegrass and tall fescue
Southern Great Plains.
in the
Increased water may not alter digestibility
because of water’s inability to significantly change the ratio of
digestible compounds and lignin. That is, there may be more of each
compound in a stjand of watered forage, yet the ratio appears to be
similar regardless of water regime.
with digestibility,
Phenology, a common correlate
was also unaffected by increased avaiable water
(Weaver and Welker in preperation), which suggest that some other
seasonal
indicator plays a major role in triggering changes in
concentration of m e t a b o l i c
compounds.
Photo-period,
another
environmental parameter which is closley associated with hormonal
fluctuations within the plant,
enzymatic
systems
parameter
involved
metabolic compounds.
(Salisbury
in
and is linked to temperature and
and
triggering
Ross
1978)
changes
might
in
be another
concentration
of
Photo-periodic control of growth stages may be
highly significant as a m e c h a n i s m
of
altering
digestibility.
Investigating the digestibility of Agropvron smithii and Bouteloua \
gracilis in which vegetative growth was maintained without flora
induction would be of interest.
Also worthy
of investigation would
28
be the physiological triggering mechanism involved in the formation of
cinnamic acid derivitives of which lignin is one (Bonner and Varner
1965).
Relative digestibility of both grasses increased after the fall
watering in 1977.
This increase suggested that after the watering,
the forage was able to respond with vegetative, regrowth,
presumably
lower in lignin concentration and therefore more digestible.
However a similar
increase was
is speculated that regrowth
not detected in the fall of 1979.
It
of Agroovron smithii and Bouteloua
gracilis did indeed occur after the fall watering in 1979, yet due to
the large amount of biomass produced in response to the water applied
in the fall of 1978, abundant standing dead material may have masked
the digestiblity of the regrowth.
This masking effect did not occur
in I 977 because no water was applied in the fall of 1976, therefore
the large amount of mature lignified forage was not present.
The fact that digestibility continued to decline from October to
November as contrasted across years for both western wheatgrass and
blue grama is of interest.
may
possibly
This decline late in the growing season
be explained
by several
lignification of the plant tissue,
f a c t o r s ; a) continued
b) transport below ground of
digestible compounds, and c) possible leaching of digestible compounds
enhanced by fall precipitation.
The possible ecological implications of increased
moisture are
29
many (Perry,
1976).
Responses of the native vegetation to additional
water varies from site to site.
At our sites digestibility of both
species were not significantly altered by short periods of water
supplementation.
Yet
inorder to alter digestibility in the long term
sufficient exposure to the altered water conditions would have to
occur to allow approprate mutations to occur (Perry 1976).
period
of
continued
water
supplementation lengthened
As the
then the
probability of natural selection for different enzmatic pathways,
photo-periodic responses and alteration of the ratio of metabolites
may increase.
Any effect on forage digestibility of long term
alteration of the local water regimes has yet to be reported.
I would hypothesis that long term alteration of the local water
regime of any native rangeland plant community would give rise to
genotypic
alteration
changes with in
of
the population,
digestibility.
One might
possibly
also expect
leading
to
that with
continued alteration of the local water regime a shift in the species
occuping the site may occur.
Therefore this shift may led to a
change in the season of highly digestible forage.
Also of interest is
the effect of supplemental water on the percent of the plant which is
greater than 40 to 50% digestible.
If water supplementation increased
the amount of the plant material which is greater than 50% digestible
then more energy and protein would be avaiable for the herbivior.
Summary
This research has documented that water applied to western
wheatgrass or blue grama at different seasons or in varying amounts
t
during the summer growing season was incapable of significantly
altering the relative digestibility of these forages „
These water
treatments were also ineffective in prolonging the decline in forage,
digestibility which commonly occurs between the vegetative and the
seed
shattering
period. ..Also
noted
was
that
season
of
water
supplementation (Fall, Spring, or Summer) did not significantly alter
the forage digestibility.
APPENDICES
32
Appendix I. Soil Descriptions.
Agropvron smithii Site
Well-drained, fine, montmorilIonitic Borollic Camborthid, with
plentiful free lime at 20cm and roots to I.Sm^ Kobar sility clay loam
which developed on alluvium lying in simple relief with a 2.% north
slope.
Bouteloua gracilis Site
Well-drained, fine loamy, frigid calcareous Ustic Torriorhent with
plentiful free lime at 75cm and roots to 105cm, Havre variant loam
developed on a high alluvial terrace with simple relief and with 1%
east slope
33
Appendix 2. Computer Program to Calculate Digestibility
and Data File.
P R O G R A M TO C A L C U L A T E % D M , O M , A S H , I V O M D,I V DM D
A = F R E S H W T . , 3 = 0 R G I NAL C R U C . W T . , C = CRU.+ D R I E D S A M PLE
D = C R U . + A S H E D S A M P L E , E = O R G . D I G . S A M P L E , F = % D W,
G = % A S H , H = O R G . G O O C H C R U C . W T . , I = DI G E S T E D S A M P . E
+GOOCH CRUC.AFTER DRYING,J=DIGESTED SAMPLE +
GOOCH C R U C . AFTER A S H I N G , K = B L A NK CO NSTA N T , O = IVOMD
M = IVDMD,N =% 0 M , L = C O D E (YR,MONTH,SPEC IE S , T R T , REP)
M O N T H - J N = I , JL = 2 , AU G = 3 , O C T , N O V = A
Y E A R , 9 = 1 9 7 9 , 8 = 1 9 7 8 , f = 1 9 7 8 ; SPECIES, I = W W H E A T G R A S S , Z = B G R A M
TREAT, 1=C0NTR0L,Z=SMM/WK,3=12MM/WK,A=FWET,S=SPWET,S=WET
INTEGER G
REAL H,I,J,K,L,M,N
D O 1 5 I I = 1 ,A I 6
READ (50,10)A,3,C,D,E,F,G,H,I,J,K,L
10
FORMAT (F5.3,X,3(FS.3,X),F6.5,X,I2,X,I2,X#
*3(F8.5,X) ,F6.5,X,I5)
F = ( ( C - B ) Z A ) *103
G = ( ( D - B ) / A )* I 03
N=(1-(G/100.0))
O = ( ( E * G /I 0 0 . O ) - ( J - H - K ) ) / ( E * C / 1 . 3 0 . 0 )
M = ( ( E * F / 1 0 Q . 0 ) - ( I-H-K)) / (E*F/1 30.0)
WR IT E ( 5 5 , 2 3 ) A , B , C , D , E , F , G , N , I , J , K , L , M , N , 0
20
FORMAT(F5.3,X,3(F6.5,X),F6.5,X,I2,X,I2,X,
*3(F8.5,X),F6.5,X,I5,X,F6.2,X,F6.2,X,F6.2)
15
CONTINUE
END
34
Appendix 2
Continued.
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35
Appendix 2.
Continued.
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34.48542
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36
Appendix 2.
A
1 7 . 0 55
1 7 .5 6 8
I Ili
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B
:89:
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Continued.
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37
Appendix 2.
A
B
1. 067 17.770
1 . 0 2 0 I 7. 980
1 . 0 0 8 I 7.671
*
17.468
Continued.
E
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18.811
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38
Appendix 2.
Continued.
B
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39
Appendix 2.
A
Continued.
B
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40
Appendix 2.
A
B
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LITERATURE CITED
' L I T E R A T U R E CITED
Albertson, F.W., and G.W. Tomanek. 1965. Vegetation changes
t
during a 30-year period in grassland communities near Hays, Kansas.
Ecology 46: 714-720.
Bokhari, U.G. 1978. Nutritional characteristics, of blue grama
herbage under the influence of added water and nitrogen. J. Range
Manage. 31: 18-22.
Bonner, J., and J.E. Varner.
Press, New York.
1965.
Plant Biochemistry.
Academic
Branson, F.A. 1956. Range forage production changes on a water
spreader in southeastern Montana. J. Range Manage. 9: 187-191.
Cogswell, C., and L.D. Kamstra. 1976. The stage of maturity and
its effect upon the chemical composition of four native range species.
J. Range Manage. 29: 469-473;'
Collins, D., arid T. Weaver.
1978. Effects of summer weather
modification (Irrigation) in Festuca idahoensis-Agropyron spicatum
grasslands. J. Range Manage. 31: 364-369.
Cook, C.W., and L.E. Harris.
1968. Nutritive value of seasonal
ranges. Utah Agr. Exp. Sta. Bull. 472.
Dehority, B.A., and R.R. Johnson, and H.R. Conrad. 1963. Digestion
of forage hemi-cellulose and pectin by rumen bacteria in-vitro and
the effect of lignification. J. Dairy Sci. 45: 508-511.
Dickerson, C.E., and J.L. Dodd.
grass prairie. Amer. Mid. Nat.
1976. Phenology pattern in short96: 367-378.
Eck, H.V., G-C. Wilson, and T. Martinez.
1981. Fescue and smooth
bromegrass: Effects of nitrogen fertilization and irrigation regimes
on quality. Agronomy
73: 453-456.
Erickerson, A. 1966. Effects of frequent clipping at different
stubble heights on western wheatgrass (Agropyron smithii).
Agronomy 58: 33-35.
r
,
43
Erickerson, D.O., W.T. Baker, and C.N. Haugse. 1978. The feeding
value of native grasses in the Sheyenne National Grasslands. North
Dakota Farm Research 36: 8-12.
Havstad, K.M. 1981.
Humphrey, R.R.
1962.
(personal communication).
Range Ecology.
Ronald Press Co. New York.
Kamstra, L.D. 1973. Seasonal changes in quality of some importnat
range grasses. J. Range Manage. 26: 289-291.
Kmastra, L.D., D.L. Schentzel, J.K. Lewis, and R.L. Eldenkin. 1968.
Maturity studies in western wheatgrass. . J. Range Manage. 21: 235239.
Launchbaugh, J., and H. Hackerott. 1969. Early spring blue grama
inflorescences from fall initiated spikes. Crop Sci. 9: 631-633.
Monson, O.W., and J.R. Oueensberry. 1958. Putting floodwaters to
work on rangelands. Montana Agr. Exp. Sta. Bull. No. 453,
Moore, J.E., and D.G. Dunham. 1978. Procedure for the Two-Stage
In-Vitro Organic Matter Digestion of Forages (Revised). Nutrition
Laboratory, Dept, of Animal Science, University of Flordia,
Gainesville, Fla.
Perry, D.A. 1976. The effects of weather modification on the Northern
Great Plains grasslands: A preliminary assesment. J. Range Manage.
29: 272-278.
Pieper, R.D., J. Montoya, and D. Grace. 1971. Site characteristics
on Pinyon-Juniper and Blue Grama ranges in Southcentral New Mexico.
New Mexico Agr. Exp. Sta. Bull. 573.
Rogler, G.A., and H.J. Haas. 1947. Range production related to soil
moisture and precipitation in the Nothern Great Plains. J. Amer.
Soc. Agron.- 39: 378-389.
44
Salisbury, F.B., and C.W. Ross. 1978. Plant Physiology, 2nd
edition. Wadsworth Publishing Co. Inc. Belmont, Calif, pp 224332.
Smika, D.E., H.J. Hass, and J.F. Power.
1965. Effects of moisture
and nitrogen fertilizer on growth and water use by native grass.
Agronomy 57: 483-486.
Tilley, J.M.A., and RiA. Terry. 1963. A two stage technique for
the in-vitro digestion of forage crops. J. Brit. Grassl. Soc. 18:
104-130.
Weaver, T.W. 1975. Effects^bf precipiation modification on Idaho
fescue medows. Proceedings 28th Ann. Meeting Soc. Range Management.
Mexico City (abstract).
Weaver, T., and J.M. Welker 1982 (manuscript in preparation).
I
0^ Agropyron smit
3 1762 00116383 9
N?78
W457
cop.2
Welker, J. M.
Relative digestibility
of Agropyron Smithii and
Bouteloua Gracilis grown
under six water regimes
✓