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RANGE
NUTRITION AND ITS
APPLICATION TO
MANAGEMENT
-by
C. Wayne Cook
COMPARATIVE
NUTRITIVE VALUES OF
FORBS, GRASSES AND SHRUBS
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The quality of a range plant is
judged primarily on palatability or how readily it is eaten by animals,
and its nutritive content with respect to its phenological development.
Some plant species are eaten only during early growth and, in special
cases, only certain portions of a particular species are consumed
during the latter stages of development. |
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During their annual life cycles, individual plant species vary materially
in nutrient content. Most forage plants are high in nutrients during early
growth. However, as they mature they lose nutrients markedly. Some plant
species mature rather rapidly and as a result decrease substantially in
nutritional value; other species mature rather slowly and consequently remain
high in nutrient content over an extended period. Still other plant species,
even after they mature, appear to cure rather well and retain comparatively
high quantities of nutrients over indefinite periods.
The
true test of the nutrient value of a forage species, or a mixture
of species, is the ability of the usable forage to meet the nutritional
require-ments for the physiological functions being performed
by the grazing animal during the various seasons of the year.

Most
physiological functions of the grazing animal are performed as
every-day functions and are generally referred to as "maintenance
requirements" which include travel, mastication, digestion,
maintaining body temperature, and growing of hair or wool.
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Of
the three forage classes, shrubby species, in most cases, contain
higher levels of carotene, lignin, phosphorus, calcium and protein
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Grasses
are higher in crude fiber, cellulose, and gross energy.
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In
most cases, forbs are intermediate between browse and grasses in
chemical content.
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The
nutrient evaluation of most range areas can be based on how much protein,
energy, phosphorus, and carotene are in the forage plants. Carotene, phosphorus,
digestible protein,and energy content are the four main nutrients that
are most important in evaluating the status of range nutrition.
The
comparative nutritive value among the forage classes (browse, forbs, grasses)
can best be discussed by measuring their apparent ability to meet the
nutritional requirements of large herbivores for the more important physiological
functions such as maintenance, gestation, growth, and lactation.
In
addition to the influence of relative preference and floristic composition
on nutritive content of range forage, there are other important factors
such as stage of growth and variable site conditions.
NUTRITIONAL
CONTENT AND STAGE OF GROWTH
For
the most part, rangelands of the West can be evaluated for nutritional
value on the basis of vitamin A (carotene), phosphorus, digestible protein,
and digestible energy. There may be some local areas where other nutrients
should be considered. During the period of initial growth, and for a time
thereafter, all forage classes are high in nutrient content, and their
nutritional contents are not materially different. However, as growth
stages advance, the nutritional differences among forage classes become
more evident. It is generally agreed that young growing animals and lactating
animals have comparable requirements when expressed on a percentage basis.
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It
is a common belief that animals grazing green plant growth on spring
and summer ranges receive adequate nutrients; however, on fall and
winter ranges where plants are dry and mature, the diets of grazing
animals are thought to be deficient in some nutrients. |
Several studies have shown that chemical composition varies with season,
largely as a result of change in the stem-to-leaf ratio and the normal maturing
process that causes a translocation of nutrients within the plant parts.
In addition to the actual decrease in chemical content of herbage with advancement
of growth, the forage likewise decreases in digestibility because of lignification
and calcification of the plant material.
CAROTENE
Browse
species decrease only slightly in carotene content from early growth to
maturity (Figure 1) because of the high carotene content in the covering
of bark on young twigs. Grasses lose their carotene rather rapidly during
advanced stages of maturity and contain almost none after they turn straw-colored
during quiescence. Carotene content places forbs in an intermediate position
between browse and grass, but forbs generally contain only a small amount
of carotene by the time the leaves are lost and the stems are dry and dormant.
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Figure
1--Average carotene content of the three forage classes compared
to phenological stage of development and requirements for lactation
and gestation.
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Browse species in all stages of growth furnish enough carotene to meet
the vitamin A requirements for even lactating or growing animals. However,
species of grass and forbs, when mature, fail to furnish even the minimum
requirements of vitamin A for gestating animals. Vitamin A has body reserves
for 90 to 120 days. Deficiencies cause abortion and night blindness.
DIGESTIBLE
PROTEIN
There
is a rather high correlation between total protein and digestible protein
in the mixed ration normally used in farmstead feeding. In range forages
the digestibility of protein may vary from 70 percent in early growth
to as low as 10 to 15 percent in the quiescent stages. It thus appears
more logical to evaluate grazing animals' diet on the basis of digestible
protein rather than on total protein unless standards have considered
the high variability in the digestibility of protein as plants mature.
Grass
species decline in digestible protein rather rapidly and generally fail
to meet the lactation requirements at about the time they come into full
anthesis (Figure 2). This is the result of a rather rapid loss of total
protein and a more rapid decrease in digestibility of protein that occurs
with advanced growth. Grasses, in general, lose 75 percent of their protein
during the period from early growth to seed formation; on the other hand,
browse lose only about 40 percent of their protein content during a similar
period. As a result, grasses that have matured are considered poor sources
of digestible protein. Shrubs, however, are considered good sources of
digestible protein during most of their active growing period; and even
after they reach full maturity they continue to meet gestation requirements.
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Figure
2--Average content of digestible protein for the three forage classes,
compared to phenological development and herbivore requirements for
lactation and gestation. |
Forbs are intermediate between browse and grass with respect to protein
content during most seasons. Most forb species fail to furnish adequate
digestible protein to meet the requirements of animal gestation after reaching
the fruiting stage.
PHOSOPHORUS
Even when mature, shrubs
are generally considered good sources of phosphorus for general animal
maintenance and gestation, unless they are deciduous. Even deciduous shrubs
are perhaps only borderline if the young twigs are readily eaten (Figure
3). Most forbs have a phosphorus content that is only slightly lower than
that of shrubs. Grasses, however, are low in phosphorus soon after they
form seed; so they are considered poor sources of phosphorus during quiescence.
Most grasses lose considerable phosphorus content when temporarily forced
into dormancy by even brief periods of drought. However, when precipitation
occurs and growth is renewed, the phosphorus content increases and lactation
requirements are again met.
| Figure
3--Average percent phosphorus for three forage classes compared to
stage of growth and herbivore requirements for lactation and gestation. |
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ENERGY
In some cases, neither
grass energy nor digestible energy in shrubs is considered a good index
to the true energy values of forage because of the high content of essential
oils, resins, or waxes that suggest high energy content; but these materials
are not available for livestock metabolism. The digestible energy values
for browse shown in figure 4 have been adjusted for species high in essential
oils so that the trends are based on realistic digestible energy values.
Shrubs are not considered good sources of energy after they reach the
phenological stage of fruit development. Thereafter, they generally fail
to meet the energy requirement for animals in gestation.
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Figure
4--Average content of digestible energy for three forage classes compared
to various stages of phenological development and requirements for
lactation and gestation. |
Grasses are generally considered
good or excellent sources of energy, primarily because of their high content
of cellulose. Even when they reach maturity, grasses seldom cease to furnish
the energy requirements for lactation. Forbs are intermediate between
shrubs and grasses in energy-furnishing constituents and, like shrubs,
they generally fail to meet the energy requirements for gestation after
reaching full maturity and dormancy.
SEASONAL
RANGES AND FORAGE CLASSES
In the intermountain area
of the West, ranges are identified with the season that apparently best
suits their forage producing qualities.
High mountain range
is used during the summer;
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foothills are used
during the spring;
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and the desert basin
areas are used primarily during winter.
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Normally, grazing animals
bear their young and are in lactation during spring and summer. During
the winter grazing period, range forage is expected to meet the gestation
requirements (Table 1). Animals may be grazed year-round in essentially
the same locality in the grassland types of the Southwest and the Great
Plains where lactation begins with new growth in the spring and continues
throughout the summer. On most grassland ranges, gestation requirements
are met by intermittent farmstead feeding periods or by giving supplements
to the grazing animals while they are on the dormant ranges.
| Table
1. Recommended nutrient requirements for cattle and sheep under range
conditions during gestation and lactation on a dry-matter basis. Nutrient
requirements are slightly higher for sheep because smaller animals
have a somewhat higher metabolic requirement per unit of body weight. |
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Percent
of ration or amount/pound of feed
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| Phase
of Production |
DP (%)
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TDN (%)
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DE
(kcal/lb)
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ME
(kcal/lb)
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P
(%)
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Carotene
(mg/lb)
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| Gestation |
4.4
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46
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830
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665
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0.17
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0.6
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| Lactation: |
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| First
8 wks |
5.4
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57
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1120
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900
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0.22
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1.6
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| Last
12 wks |
4.5
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49
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880
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700
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0.20
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1.6
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DP
represents digestible protein.
TDN
represents total digestible nutrients. DE represents digestible
energy. Both are calculated by deducting allowance for high ether
extract in browse in the diet.
ME
represents metabolizable energy.
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Spring Ranges
The early growth of forage
plants on spring ranges results in a high nutritive content; but as discussed
previously, the percentage of nutrients decreases as the growth stages
advance. Cool-weather grasses start growth early but lose nutrients rather
rapidly, even during active growth. Warm-weather grasses start growth
later in the season than cool-weather grasses, but the former retain their
nutrients much better as the season advances.
Generally, all forage classes furnish rather large quantities of
carotene on spring range, but some grasses and forbs may fail to furnish
adequate quantities of digestible protein for lactating animals when
plants reach late anthesis in late spring. Grasses may not meet the
phosphorus requirement for lactating animals during late growth stages
during spring grazing, but browse and forbs generally do not become
deficient until at least full maturitiy is reached late in the spring
or summer.
Both forbs and shrubs may become
decidedly deficient in energy-furnishing constituents for lactating animals
when plants approach maturity during late spring, but grasses continue
to furnish adequate energy for lactating animals during the entire spring
grazing season and even later into summer and fall. Therefore, a stand
of mixed vegetation that includes all forage classes generally meets lactating
requirements for livestock throughout the entire spring and summer grazing
seasons.
Summer
Ranges
On mountain ranges where
mountain brush, aspen, and coniferous types prevail, the vegetation is
still in the initial growth stages even by late June or the first of July.
Whereas, on the grassland plains used for summer grazing, many forage
species are in rather advanced stages of growth by July 1. Intermittent
precipitation, when it falls on the grassland plains area, produces renewed
growth and thus increases the nutritional value. In both the mountainous
areas and the grassland plains, drought periods may produce temporary
dormancy in herbage and consequently produce reduced percentages of nutrient
content. In some cases, serious nutrition deficiencies may occur.
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either mountain or grassland summer ranges, the plants may reach
full maturity late in the season, and nutrient requirements may
become borderline for growing and lactating animals. |
As discussed previously,
each of the three forage classes displays characteristic changes in nutrient
content as advanced stages of growth occur. Late in the summer, grasses
would be expected to be low in protein and phosphorus, but they would
be low in energy. Forbs, as previously stated, would be intermediate between
grasses and shrubs in these respects. During the summer grazing season,
grasses may lose considerable quantities of their protein content and
increase decidedly in lignin and cellulose. Protein content of forbs and
shrubs generally decreases only slightly during the summer, while lignin
and cellulose content increase only moderately as the season advances.
Shrubs and
forbs furnish ample protein and phosphorus for growth and lactation until
(at least) late in the summer grazing season or even into the fall unless
drought conditions force plants to become dry and dormant before they
normally complete their annual cycle. However, a diet composed largely
of forbs and browse late in the summer might be deficient in the constituents
that supply energy. Most grasses would furnish adequate energy for growing
and lactating animals even late into the summer grazing season, but grasses
would be deficient in both protein and phosphorus when they reached the
advanced stages of growth. All three forage classes are comparatively
high in carotene (vitamin A) during the entire summer grazing season unless
the herbage becomes extremely dry as a result of an extended drought period.
Seasonal changes in nutrient content
on summer ranges show generally that phosphorus, protein, gross energy,
and carotene decrease in all three forage classes as the season advances.
Grasses change the most as to nutrient content: shrubs change the least,
except for changes in digestible energy. As would be expected, shrubs
have the highest protein and phosphorus content throughout the summer
grazing season, and grasses have the highest energy content. These changes
that occur with advancement of the summer grazing season are affected
by changes both in the stem-leaf ratio and in chemical content of the
plant parts themselves. Leaves for all forage classes are higher in ether
extract, protein, phosphorus, and calcium; and stems are higher in lignin
and cellulose. The leaves of forage plants are also much higher in digestibility
because of less fiber content compared to stems. Thus, leafiness leads
to higher nutrition among range plants.
Thus, it is concluded that summer ranges,
including both warm and cool weather grasses and more than one forage
class, more nearly meet growth and lactation requirements than ranges
on which there is a restricted diversity in forage species. Moving up
in elevation on mountain rangelands as the summer season advances provides
an increase in nutrient content because of delayed plant development.
Winter
Ranges
During winter on desert
ranges, shrubs generally meet the gestation requirements for livestock
regarding protein and carotene, but they are borderline in phosphorus
and are decidedly low in energy for animal metabolism. During winter,
the grasses are decidedly deficient in protein, carotene, and phosphorus
but are good sources of energy. The broad general class of shrubs on desert
ranges can be divided into two groups. The first group would include species
that possess a woody base but, for the most part, have herbaceous stem
growth. These are known as "suffrutescent" (half shrub) species.
The second group includes the true shrubs, or frutescent species. The
species of the second group include wood stemmed plsnts of low stature.
As might be expected, the suffrutescent
species are intermediate between shrubs (frutescent species) and grasses
in nutrient content (Table 2). The suffrutescent species are slightly
lower than the shrubs in these respects. In the suffrutescent species,
energy-furnishing constituents are somewhat lower than in grasses but
these species provide somewhat higher energy sources than shrubs. Again,
it is suggested that a mix of forage classes in the diet is conducive
to a balanced nutritional diet for grazing animals on desert ranges during
the winter.
Table
2--Average chemical content for three shrubs, two half-shrubs (suffrutescent),
and two grass species harvested during four seasons at three intensities
during 1968 (the sixth year of treatment).
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| Forage
class |
Protein
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Lignin
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Gross
Energy
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Phosphorus
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Percent
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Percent
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Mcal/kg
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Percent
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| Shrubs |
11.2
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12.7
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3.85
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0.12
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| Half
shrubs |
10.1
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9.7
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3.90
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.10
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| Grass |
6.2
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5.7
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4.11
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.09
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Forage
class - shrubs were big sagebrush, shadscale and black sagebrush;
half shrub (suffrutescent) species were winter fat and Nuttal's
saltbush; and grasses were Indian ricegrass and squirreltail grass.
Gross
energy - adjusted for essential oils.
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Sandstone and limestone
breaks in the Great Plains are often excellent winter ranges because of
the grass-shrub mixtures. Late in the winter when grass is the dominant
forage, carotene will become deficient in the diets of grazing animals,
but most herbivores store vitamin A in the liver as a reserve for 60-90
days.
During spring, while they
are growing, desert shrubs are as high in nutrient content as mountainous
shrubby species during comparable phenological growth stages. This is
also true for grasses and forbs, but forbs are sparse on most desert ranges
and, therefore, are not significant in the diet of grazing animals. Even
though desert plants are high in nutrients during growth, they can withstand
only about one-half the degree of defoliation that is tolerated by mountainous
and plains plants during growth.
SUPPLEMENTING NATIVE
GRASS RANGES
Predominantly grass ranges
such as short grass or mixed grass are considered deficient in phosphorus,
protein and vitamin A when the plants approach hard seed formation or
maturity (See figures 1, 2 and 3). Therefore grass ranges are generally
believed to be deficient in these three nutrients in late summer, fall
and winter and should be supplemented somehow.
Supplements
Planting
agriculture farm lands to plants that can be used to supplement rangelands
during the period that the native range forages are deficient in nutrients
is useful. This practice, in most cases, is a real money saver for supplementing
the diet of grazing animals. Examples of this are using grazing sorghums
in late summer or fall to maintain or increase the gains of steers that
will enter feed lots. Similarly, we plant domestic wheat or rye to be
grazed in the late spring to enhance or extend the period of normal spring
gains by steers that are to be placed in feed lots.
These are
all examples for obtaining gain from grazing animals in the schemes for
obtaining cheaper gain compared to the feed lot gains that would otherwise
be employed prior to full feeding for desired finished fattening. Final
finishing products for sale are shortened and total costs are materially
reduced.
Recently
we have discovered that our normal range forages from grass ranges are
decidedly deficient in nutrients in late summer, fall, and winter. These
deficiencies in grass range forage begin in late summer and continue to
decrease in nutrients as the grazing forages mature and lose their ability
to meet nutritional requirements.
During the
last few years, there has been a move to add a special plant species to
various portions of the native range by seeding in widely spaced contour
furrows or by broadcasting seeds on the range here and there; then covering
the seeds by dragging a spike-toothed harrow over the area. Only a relatively
small portion of the entire range is needed,
These methods have been used in the grassland
areas throughout the Plains states by interseeding portions of their large
grassland areas to fourwing saltbush (atriplex canescens) so that
this palatable, persistent-leaved shrub can be used to supplement their
native range grasslands. This practice came about largely because suitable
commercial supplements of phosphorus, vitamin A, and protein were costing
well over $100 per ton.
NUTRITIVE
VALUE AS AFFECTED BY SITE
Site
conditions are important because they influence the growth characteristics
of range plants and thus indirectly affect their nutritive value. Sites
also indirectly affect the chemical content of plants and plant parts
through soil and plant development, water runoff, intensity of shade,
and other environmental factors.
Shrubs and
forbs, when they approach maturity, are considerably more leafy on less
favorable growing sites. Grasses in advanced stages of growth are, likewise,
more leafy on poorer sites than on more favorable sites, but differences
between sites are less conspicuous for grasses than for either shrubs
or forbs. As a result, plants on unfavorable sites are more palatable,
more digestible and, therefore, are more nutritious than plants on the
unfavorable sites.
Nutrient
content of plants on poor sites during advanced growth stages are generally
considered more nutritious because of the finer material and the presence
of more leafy material compared to stems. The differences in stem-to-leaf
ratios would, to a large degree, account for chemical differences between
plants growing on favorable and unfavorable sites since leaves are higher
in ether extract, protein, ash, calcium, phosphorus, and nitrogen-free-extract;
whereas stems are higher in lignin, crude fiber and cellulose.

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