Escolar Documentos
Profissional Documentos
Cultura Documentos
Floral induction marks the transition of the apical meristem from a vegetative to
a reproductive status (Sharman 1945, Etter 1951, Langer 1972). Floral induction
occurs in response to a photoperiodic stimulus (i.e., day length) following a
sufficient juvenile growth period. At the time of floral induction, both leaf
primordia and axillary buds are rapidly differentiated producing a double ridge
appearance on the apical meristem. Spiklet primordia differentiate from the
axillary buds while further development of leaf primordia is suppressed halting
additional vegetative development. Vegetative growth can only occur from
immature intercalary meristems of existing phytomers or from previously
differentiated axillary buds in reproductive tillers.
top
Tillers. The accumulation of successive phytomers differentiated from a single
apical meristem defines the tiller (Etter 1951, Hyder 1972, Briske 1986). Tillers
are initiated from the axillary buds of ontogenetically, older parental tillers (Fig.
4.3). Following a juvenile period of development, fifth-leaf stage in timothy
(Langer 1956), tillers are potentially capable of initiating additional tillers
from axillary buds differentiated with each phytomer. The largest, but
ontogenetically youngest axillary buds, develop to form tillers in crested and
bluebunch wheatgrass (Mueller and Richards 1986). These observations support
the contention that axillary buds may possess a relatively brief longevity
following their development (Hyder 1974, Dahl and Hyder 1977). However, no
evidence of bud senescence was observed in either of the wheatgrass species
even though bud growth was arrested at about the time the associated leaf within
the phytomer matured (Mueller and Richards 1986).
Morphological variation of individual tillers is largely a consequence of the
number and length of phytomers comprising the tiller. Variation in tiller
architecture among tall, mid- and short grasses does not originate from a major
deviation in the pattern of developmental morphology, but rather results from a
variable number and/or size of phytomers determining cumulative tiller height.
Internode elongation increases phytomer size and is most frequently associated
with reproductive tiller development, but may also occur in nonreproductive
tillers in a small number of species. Inculmed vegetative tillers, the apical
meristem is elevated above the soil surface by internode elongation while in a
vegetative condition. Culm elongation originates from the activity of intercalary
meristems located at the base of the several uppermost internodes. In
reproductive tillers, the inflorescence and several uppermost leaves are elevated
above the soil surface presumably to facilitate wind pollination. The
developmental morphology of reproductive tillers is similar to vegetative tillers
prior to floral induction (Hyder 1974).
Figure 4.3
top
Plants. The spatial arrangement of tillers within the grass plant, in addition to
morphological variation within individual tillers, is a major determinant of
architectural variation within the grass growth-form (e.g., bunchgrasses versus
sodgrasses). Spatial arrangement of tillers within the plant is dependent upon the
pattern of tiller development. Intravaginal tiller development within the
subtending leaf sheath results in a compact spatial arrangement of tillers defining
the bunchgrass (caespitose or tussock) growth-form (White 1979, Briske
1986, Fig. 4.4). Contrastingly, extravaginal tiller development proceeds
laterally through the subtending leaf sheath contributing to greater inter-tiller
distance and tiller angles within the plant. Extravaginal tiller development is a
prerequisite to the formation of the sodgrass (creeping or spreading) growthform which may be further accentuated by the development of rhizomes and
stolons. These modified, horizontal tillers further increase inter-tiller distance
within the plant depending on whether they are determinate or indeterminate in
growth. The apical meristem of determinate rhizomes eventually emerge from
the soil to form a tiller while the apical meristem of indeterminate
rhizomes continue to grow parallel to the soil surface with individual tillers
potentially developing from axillary buds located at the nodes. Stolon growth
generally displays the indeterminate growth pattern (Hyder 1974).
The mechanisms determining whether a tiller, rhizome or stolon develop from an
axillary bud are not clearly understood. These shoot-types are initiated from a
finite number of axillary buds which may partially explain the observed
seasonality of tiller and rhizome recruitment (i.e., rapid tiller recruitment may
potentially limit rhizome recruitment). High nitrogen availability, high
temperatures and short photoperiods promote tiller development to a greater
extent than rhizome development in quackgrass (McIntyre 1967). Bermudagrass
stolons exposed to a high ratio of red:far-red radiation displayed an upward
curvature, increased leaf and internode elongation and lower carbohydrate
concentrations than stolons grown in darkness or in radiation with a low red:farred ratio (Willemoes et al. 1987). These data suggest that phytochrome (a
proteinaceous pigment sensitive to specific wavelengths) may regulate the
differentiation of stolons, rhizomes or tillers by affecting the distribution of
photosynthetic products within the plant. Radiation quality has also been
implicated in the regulation of tiller recruitment in several grasses (Casal et al.
1986, Kasperbauer and Karlen 1986).
Figure 4.4
top
Root Systems. Grasses produce two distinct root systems during their
developmental history. The initial root system, referred to as the seminal root
system, develops rapidly from the embryo upon seed germination (Langer 1972,
Hyder 1974). Although the seminal root system is essential to the initial
development of grass seedlings, it is relatively short-lived, generally surviving no
longer than the growing season in which it originated. The adventitious nodal
root system consists of whorls of roots originating from nodes along the base of
the tiller forming the permanent root system of the grass plant. Adventitious root
longevities vary from 1 to 4 years among species (Weaver and Zink 1946,
Troughton 1981).
Adventitious roots differ from seminal roots in both diameter and mass per unit
length. The large diameter of adventitious roots is associated with a greater cross
sectional area of xylem to enhance water transport to the shoot system (Wilson et
al. 1976). The large root mass per unit surface area may explain why adventitious
roots are not initiated until seedlings produce sufficient leaf area and
photosynthetic capacity to support their development 1 to 3 weeks following
seedling emergence (Wilson and Briske 1979). In the case of vegetative tiller
development a seminal root system is not initiated. Juvenile tillers are supported
by parental tillers until the adventitious root system develops (Welker et al.
1987). Adventitious root development is initiated at approximately the third to
fourth leaf stage in little bluestem (Carman and Briske 1982).
top
Tiller Demography
Tillering. The perenniality and sustained productivity of grasses are conferred by
the successive production of relatively short-lived tillers (Langer 1963,
Tomlinson 1974). Successive tiller recruitment produces a series of connected
tiller generations referred to as tiller hierarchies or families (Langer 1963). The
number of generations within a hierarchy is determined by the rate of tiller
recruitment and tiller longevity as influenced by genetic and environmental
constraints. The number of tillers per hierarchy and number of hierarchies per
plant define the size and architectural configuration of the plant. With increasing
plant size and age, these tiller hierarchies become separated as the initial tiller
generations die and decompose (Gatsuk et al. 1980). Each plant fragment is
capable of survival and may continue tiller development as previously described.
The hollow crown phenomenon characteristic of many perennial grasses is very
likely a natural consequence of the developmental morphology of grasses and not
Figure 4.5
Figure 4.6
top
Tiller Longevity. Tiller longevity in temperate perennial grasses is approximately
1 year and does not exceed 2 years (Langer 1956, Robson 1968, Butler and
Briske 1988, Briske and Butler 1989). Longevity is directly influenced by season
of tiller recruitment and phenological development. Tillers recruited early in the
growing season will have the greatest probability of becoming reproductive and
terminating growth at the end of the season in which they were recruited. Tillers
initiated later in the season apparently do not surpass the juvenile growth
requirements necessary to respond to the long-day photoperiodic stimulus.
Consequently, these tillers may overwinter in a vegetative stage and resume
growth the subsequent spring. Tiller growth, including dry weight, leaf number
and seed yield, is greatest in tillers initiated in the latter portion of the previous
season or early in the season of reproductive development because they
experience a longer period for growth and development (Langer 1956).
top
Leaf Demography
Leaf longevity also displays seasonal patterns of recruitment and mortality (Vine
1983, Chapman et al. 1983, 1984). Leaves initiated when growing conditions are
most favorable, spring and early summer in temperate environments, have shorter
longevities than those initiated during periods of less favorable environmental
conditions. Leaves of perennial ryegrass and browntop exhibited mean
longevities of 60 - 70 days when initiated in the spring and summer compared to
70 - 105 days when initiated in the fall and winter (Chapman et al. 1984). Leaf
longevities of less than 90 days during favorable growing conditions indicate that
grazing must closely follow leaf initiation to optimize the harvest of live leaves.
The environmental conditions experienced by emerging leaves may program
their subsequent development to a larger extent than the conditions during growth
or maturity.
Synchronous leaf initiation and senescence maintains a relatively constant leaf
number per tiller throughout much of their developmental history. Generally, a
tiller possesses an emerging leaf, immature leaf, mature leaf and senescing leaf
(Anslow 1966, Chapman et al. 1984). The net difference between leaf initiation
and senescence represents the number of live leaves per tiller. Leaf demography,
by determining the amount of live leaf area per tiller, influences both the
potential photosynthetic capacity of the tiller and the amount of leaf biomass
available for consumption by herbivores. Leaf and tiller demography collectively
determine the rate of biomass turnover (i.e., production versus senescence) within
the community.
top
Benefits of Vegetative Growth
Grasses exhibit vegetative growth or reproduction by the successive recruitment
of tillers from previous generations. Each tiller establishes a shoot and root
system (adventitious) to acquire resources from the environment. Vegetative
growth may confer several ecological advantages originating from resource
allocation between and among tillers within individual plants (Pitelka and
Ashman 1985). The capacity for continuous tiller replacement and site
occupation based upon parental support of juvenile tillers is perhaps the most
significant ecological benefit. Resource allocation from parental to juvenile tillers
confers a greater likelihood of establishment and survival in comparison with
seedlings which must become established from energy and nutrient reserves
available within the endosperm (Tripathi and Harper 1973).
Survival and growth of stressed tillers are also enhanced by resource import from
associated non-stressed tillers within a plant (Gifford and Marshall 1973, Ong
and Marshall 1979, Welker et al. 1987). Root growth, as estimated by both total
length and numbers of roots, of recently recruited juvenile tillers progressively
decreased as the juvenile tiller, the parental tiller or the parental tiller and all
remaining tillers within little bluestem plants were defoliated (Carman and Briske
1982). Greater growth rates were observed in tall fescue plants following
defoliation as the percentage of undefoliated tillers within the plant increased
(Matches 1966). Both observations support the conclusion that defoliated tillers
were deriving support from associated, nondefoliated tillers.
Vegetative growth theoretically confers plants with potential immortality.
Individual plants of hard fescue have been estimated to be greater than 1000
years old (Harberd 1962). However, the few age estimates available for North
American perennial grasses indicate that their life spans are relatively short.
Estimates of individual plant longevities on the Jornada Experimental Range in
New Mexico, including tobosa grass, black grama and red threeawn, indicate that
maximum plant longevity does not exceed 30 years (Wright and Van Dyne 1976).
These estimates of plant longevity are corroborated by the work of West et. al.,
(1979) on the U.S. Sheep Station in Idaho and Canfield (1957) on the Santa Rita
Experimental Range in Arizona.
top
Grazing Resistance
Grazing resistance is an ambiguous term used to describe the relative ability of
plants to survive grazing. However, strategies to cope with grazing vary greatly
in form and expression among plant species. Additional insight can be gained by
organizing grazing resistance into a tolerance and avoidance component (StuartHill and Mentis 1982, Briske 1986, Fig. 4.7). Avoidance mechanisms reduce the
probability and severity of plant defoliation (i.e., escape mechanisms),
while tolerance mechanisms facilitate growth following defoliation (i.e.,
mechanisms of rapid leaf replacement). The ability of a species to survive
grazing undoubtedly results from a combination of these two components, but in
certain species and under specific environmental conditions, one component may
predominate over another.
Figure 4.7
Grazing resistance within ecological plants groups may be generally ranked as
follows: herbaceous monocots > herbaceous dicots > deciduous shrubs and trees
> evergreen shrubs and trees (Archer and Tieszen 1986). This ranking is based
upon both morphological and physiological considerations. Apical and
intercalary meristems within monocots are less vulnerable to large herbivores
because of their basal location within the plant. Meristems are located at terminal
and lateral positions of shoots in dicots increasing their susceptibility to large
herbivores. Many woody plants, evergreen species most notably, possess slow
growth rates and low rates of resource acquisition which require that individual
leaves be retained for long periods (Chapin 1980). Consequently, these species
are less efficient in replacing photosynthetic surfaces removed by herbivores and
frequently rely on avoidance mechanisms rather than tolerance mechanisms to
cope with grazing.
top
Grazing Avoidance
Mechanical Mechanisms. Avoidance mechanisms primarily influence plant
accessibility and palatability to specific herbivores. At the individual tiller level
of organization, the probability and severity of defoliation may be reduced by a
number of avoidance mechanisms originating from a variety of morphological
parameters (Fig 4.8). Tissue accessibility is primarily a function of tissue
proximity to the soil surface as influenced by the length and angle of leaves and
increase during the growing season (Caldwell et al. 1983). In addition, selective
grazing may increase the proportion of water utilized by the less preferred
species within the community.
top
Root growth and function. Root growth and function is dependent upon energy
provided by photosynthesis. Consequently, the suppression of root growth is
generally proportional to the intensity and frequency of above-ground defoliation
(Crider 1955, Cook et al. 1958, Youngner 1972). A single defoliation removing
50% or more of the shoot volume retarded root growth for 6 - 18 days in seven of
eight perennial grasses investigated (Crider 1955). A single defoliation removing
80 and 90% of the shoot volume stopped root growth for 12 and 17 days,
respectively. Multiple defoliations detrimentally influenced root growth to a
greater extent than single defoliations. The initial removal of 70% of the shoot
volume followed by three subsequent clippings per week stopped root growth for
the entire 33 day investigation in all three species subjected to multiple
defoliations. Cessation of root growth has been observed to occur within hours of
defoliation (Davidson and Milthorpe 1966, Hodgkinson and Baas Becking 1977).
Root growth cessation affects both lateral and vertical development of root
systems (Schuster 1964, Smoliak et al. 1972) as well as detrimentally influencing
root initiation, diameter, branching and total production (Biswell and Weaver
1933, Jameson 1963, Evans 1973, Carman and Briske 1982, Richards 1984).
Root mortality has also been observed following defoliation (Weaver and Zink
1946, Hodgkinson and Baas Becking 1977, Troughton 1981). These detrimental
responses collectively serve to reduce the total absorptive surfaces and soil
volume explored for water and nutrients.
The capacity for nutrient absorption per unit length in temperate, perennial
grasses parallels the growth responses following defoliation. Phosphorus
absorption, root elongation rate and respiration rate remained suppressed for an
8-day observation period following defoliation of orchardgrass to a height of 2.5
cm (Davidson and Milthorpe 1966). Similar responses of root growth and
function to defoliation originate from the dependence of both processes on
energy produced in plant respiration (Caldwell et al. 1987). Respiration, in turn,
functions upon substrate produced in photosynthesis.
top
Apical dominance. Apical dominance within the annual grasses, teosinte and
barley, was initially described by Leopold (1949) as the production of auxin
within the apical meristem which suppressed axillary bud expansion. This line of
experimentation was apparently based on the work of Thimann and Skoog (1933)
who had established that auxin controlled branching in dicots. This single
investigation (Leopold 1949) has largely shaped our perception of how the
tillering process is regulated in grasses, but has been criticized as being less than
definitive from an experimental perspective (Williams and Langer 1975).
Aspinall (1961) later forwarded the nutritive theory which suggested that interorgan competition for nutrients inhibited axillary bud development. This theory
was deemed untenable based on the relatively small metabolic demand presented
by both the apical meristem and axillary buds. These two theories were
eventually combined and extended into the nutrient diversion theory of apical
dominance (Jewiss 1972, Hillman 1984). This theory proposed that growth
regulators control both the supply of photosynthetic products received by axillary
buds and the rate of cellular division and expansion within the buds. The
discovery that cytokinins and potentially other growth regulators are involved in
the regulation of bud growth marked a significant advance in the understanding
of apical dominance (Phillips 1975). The principle role of auxin produced in the
leaf primordia of apical meristems is to limit the availability or utilization of
cytokinin within axillary buds thereby inhibiting growth. Although the complete
mechanism of apical dominance is not thoroughly understood, the direct
suppression of axillary bud growth by auxin is no longer an accurate
interpretation of the phenomena.
It would appear unduly restrictive to presume that only the removal of apical
meristems by grazing could serve as an environmental cue to induce tillering.
How would tillering occur in plant populations subjected to limited grazing?
Tiller recruitment has been observed to occur in response to grazing even though
apical meristems were insufficiently elevated to be removed by livestock (Butler
and Briske 1988). Conversely, removal of apical meristems from tillers of crested
and bluebunch wheatgrass did not always result in accelerated tiller recruitment
(Olson and Richards 1988b, Richards et al. 1988). In contrast to the temperate
species, lateral bud growth was stimulated by both apical meristem and canopy
removal in three tropical grasses. Expanding leaves and either the inflorescence
or elongating culm were observed to be the source of apical dominance in
vegetative and reproductive ryegrass tillers, respectively (Laidlaw and Berrie
1974). These conflicting observations attest to the complexity associated with the
regulation of tiller recruitment in perennial grasses (Youngner 1972).
Light quality has been implicated in the control of axillary bud expansion in
several grass species (Deregibus et al. 1985, Casal et al. 1985, 1986, Kasperbauer
and Karlen 1986). This photomorphogenetic response is presumably mediated by
phytochrome, as is flowering and branching in many dicotyledonous species. A
decrease in the ratio of red:far-red radiation associated with increasing canopy
physiological processes at the tiller and plant levels to enhance growth following
grazing. Both components contribute to grazing resistance, but the relative
magnitude and associated cost of each component are poorly understood.
Grazing management modifies competitive interactions by influencing the
frequency, intensity and seasonality of plant defoliation (Fig. 4.13). Plant species
grazed less frequently and intensively, or with a greater capacity to grow
following defoliation, display greater leaf areas for photosynthesis and attain a
competitive advantage. Grazing-induced modifications in competitive
interactions are eventually expressed as modifications in plant and population
structure. A decrease in total basal area, plant density or tiller density of a given
species is ultimately manifested in a relative reduction in resource acquisition
within the community. Shifts in species composition subsequently alter the
quantity, quality and variability of plant production by modifying the amount and
pattern of energy flow through the ecosystem (see Chapter 1).
Figure 4.13
Vegetation response to grazing may be investigated at one or more levels within
the hierarchical organization of grasslands (e.g., tiller, plant, population or
community). For example, plant productivity may be reduced by a decrease in
individual tiller weight, tiller number per plant, or plant density in response to
grazing. Consequently, insight into mechanisms occurring at higher hierarchical
levels (e.g., community) requires that processes at lower hierarchical levels (e.g.,
population, plant and tiller) be investigated (Archer and Tieszen 1986, Brown
and Allen 1989, see Chapter 5). Hierarchical levels of vegetation organization
may respond in a comparable manner to affect vegetation dynamics, but
frequently additional complexity is encountered by the occurrence of opposing
responses between or among levels. Grazing has been observed to increase tiller
density, but concomitantly decrease individual tiller weight (Jones et al. 1982) or
increase plant density while reducing basal area per plant (Butler and Briske
1988). It is essential that several hierarchical levels be considered when
evaluating vegetation responses to grazing to avoid incomplete or erroneous
conclusions.
Research oriented at the population level of vegetation organization possesses the
potential for integrating the divergent sources of information available from
individual plant and community studies. These two research perspectives have
not been effectively unified into an information base for vegetation management
in grazed systems. Community level investigations describe species composition
shifts and biomass dynamics, but do not yield insights into mechanistic causeeffect relationships. Conversely, reductionist investigations at the individual plant
level yield mechanistic insights, but are frequently too narrow in scope to
identify interactions and properties of systems at levels of organization suitable
for vegetation management. A major limitation to the extrapolation of plant level
studies is the minimal amount of information concerning competitive interactions
and population ecology of dominant plant species.