GRAZING MANAGEMENT An Ecological Perspective Edited by Rodney K. Heitschmidt and Jerry W. Stuth TIMBER PRESSCHAPTER 1 AN ECOLOGICAL PERSPECTIVE D. D. Briske and R. K. Heitschmidt INTRODUCTION Grazing or herbivory is the process by which animals consume plants to acquire fnergy and nutrients. Grazing management involves the regulation of this consump. five process by humans, primarily through the manipulation of livestock, to mest specific, predetermined production goals. Both the grazing process and associated Tianagerial activities occur within ecological systems and are therefore subject to an identical set of ecological principles which govern system function. These ecological Principles impose an upper limit on animal production which cannot be overcome by management. The fact that both the grazing process and efforts to manage it are influenced by a common set of ecological principles justifies the evaluation of grazing management in an ecological context, Humankind has historically fostered and relied upon livestock grazing for a sub- Santial portion of its livelihood because itis the only process capable of converting the energy in grassland vegetation into an energy source directly consumable by humans, Biochemical constraints determine that herbivores, function as “energy brokers” between the solar energy captured by plants in the photosynthetic process and its subsequent use by humans (Southwood 1985). The inability of humans to Girectly derive caloric value from the 19 billion metric tons of vegetation produced annually in tropical and temperate grasslands and savannas (24 millions km?; Leith 1978) Provides the ultimate justification for evaluating grazing as an ecological process. Ecological processes associated with grazing have probably not changed appre- Ciably since the initial appearance of grasses and grazers in the fossil record some 45 illion years ago (Stebbins 1981). However, a rapidly expanding human population, escalating degradation of natural resources, and increasing socio-economic pressures have all increased the complexity associated with the management of grazed systems Hundreds of experiments have been conducted and thousands of pages written addressing the ecological processes and plant and animal responses within grazed systems. Unfortunately, little attention has been directed toward an interpretative synthesis of this information. The diverse subject matter encompassing several disciplines (e.g, ecology, animal science, hydrology, economics, and systems science), and the difficultly associated with identifying an organizational scheme capable of encompassing this large body of information, have undoubtedly con- tributed to the lack of subject matter synthesis, The absence of a unified conceptual famework has impeded both the study and the management of grazed systems. The objective of this chapter is to identify and evaluate the fundamentalecological systems; and conclude with a discussion of the ability of humans to regu- late the ecological processes affecting plant and animal production within grazed systems. The topic is treated in a broad, conceptual manner to provide the organiz: tional framework necessary to evaluate the relative relationships among specific com- ponents and processes within grazed systems. Specific subject matter areas are ‘reated in greater detail in subsequent chapters. owes wyatt, eeuitL ECOLOGICAL SYSTEMS Structure of Ecological Systems Ecological systems or ecosystems are defined as assemblages of living organisms in association with their physical and chemical environment. The ecosystem concept is intended to demonstrate the interrelationship or interdependence among the various components within a system, rather than to delineate a specific set of organisms within a geographic area (Odum 1971; Begon et al. 1986). Consequently, ecosystems are arbitrarily defined depending upon the interest of the investigator. The living (biotic) component of ecological systems is classified according to the strategy organisms use to acquire energy and nutrients from the nonliving (abiotic) component. The two most basic strategies are autotrophic (self-nourishing), and heterotrophic (other-nourishing) (Odum 1971; Begon et al. 1986). Autotrophs acquire energy from solar radiation by photosynthesis, while heterotrophs acquire energy by ingesting other organisms. Autotrophs or producers include all species of green plants, while heterotrophs or consumers encompass all animal species including microorganisms (Fig. 1.1). The abiotic component defines the physical and chemical components of the system. The ultimate energy source, solar energy, and vaw materials (e.g, CO,, HzO and nutrients) necessary to convert solar energy into chemical energy are also present within the abiotic component. Structure of Ecological Systems Abiotic Component Rodiation soils Climote Geography ‘Atmosphere Fire Biotic Component Teetiry Consumers pe Figure 1.1. Generalized description of the structure of ecological systgms. The abiotic (a: living) component comprises the physical and chemical environment of the biotic (living) component.An Ecological Perspective 13 Function of Ecological Systems Energy Flow Energy flow within ecological systems may be viewed in terms of an economic analogy (Gosz et al. 1978). Economics, like ecology, is concerned with the movement of valuable commodities through a series of producers and consumers. A viable ecological system depends on the flow of energy just as a viable economy depends on the exchange of currency. An analysis of energy flow provides a balance sheet for monitoring energy inputs and outputs within a system not unlike a ledger of credits and debits. The magnitude and efficiency of energy flow between various feeding levels within systems can also be evaluated. The initial capture of solar radia- tion by vegetation, the efficiency of vegetation utilization by herbivores, and the effi- ciency with which ingested energy. is converted into animal growth comprise the major energy transfer processes in grazed systems. Equally important, but less obvious, is the influence energy flow exerts on the managerial components of systems including resource availability, supply:demand ratios, and price-market structure. Solar energy is initially converted into chemical energy by photosynthesis within the chlorophyll-containing cells of planfs. The energy captured within plants is subse- quently transferred to one of two general categories of heterotrophic organisms (Fig. 1.2). In the absence of herbivores, energy is transferred ‘directly into litter following plant senescence. A series of microorganisms, primarily bacteria and fungi within the soil (decomposers), utilize this organic matter as an energy source eventually releasing heat energy in association with microbial respiration. This pattern of energy flow defines the detrital food chain (Golley 1960; Odum 1971). In the presence of herbivores, a portion of the energy initially captured by plants is consumed and con- verted into animal tissue. Herbivores, in turn, may be ingested by other consumers at higher feeding levels (ie., carnivores and humans). Heat energy is released by consumer respiration at each feeding level, This pattern of energy flow through the system defines the grazing food chain. Energy is transferred from the grazing to the detrital food chain in the form of feces and animal tissue following death. ‘Two thermodynamic laws govern the flow of energy within ecological systems (Golley 1960; Odum 1971). The first law states that energy can be transformed from one form to another (e.g, conversion of solar energy to chemical energy by photo- synthesis), but cannot be created or destroyed. The second law establishes thet Producer Respiration Consumer Respiration Consumers Decomposers X Decomposer Respiration Figure 1.2. Simplified illustration of energy flow through ecological systems. Solar energy is initially captured by primary producers and transferred through at least one consumer feeding level to form the grazing food chain, or directly into the decom- poser compartment to form the detrital food chain (after Whittaker 1972).44 Uv, priske ana & AR. menstiumet energy transformation processes are not 100% efficient. These laws dictate that alarge proportion of the chemical energy, approximately 90%, transferred between feeding whic) levels within a system is converted to heat energy of limited value to the biotic portion of the system. The energy “loss” between feeding levels results from en ineffigent transfer of organic matter (e.g., gaseous, urinary, and fecal losses) and the cnergy required for internal maintenance of organisms (ie, maintenance energy). For example, only a portion of the solar energy converted into chemical energy by photosynthesis is realized as growth because a portion is utilized in respiration Similarly, animals utilize a large portion of the total energy ingested for basal metabolism, thereby diminishing the amount of energy available for growth or trans fer to subsequent feeding levels within the system (see Chapter 2). The capacity of ecological systems to produce biomass would initially appear limitless given the large, continuous supply of solar energy. However, above-ground primary productivity (plant growth/area/time) is less than 1000 kg/ha/yr in many rasslands (Sala etal, 1988) Primary productivity is limited by two general categories of ecological constraints, The first constraint involves the quality of solar radiation gvailable at the earth’s surface. Only about 45% of the solar energy is within the appropriate region of the spectrum to be effective in photosynthesis (Gose etal. 1978; Begon etal. 1986). The remaining 55% of the spectrum is primarily composed of long- wave thermal energy unavailable for conversion into chemical energy. However, this energy affects ecological systems, in that it is absorbed as heat energy by the atmosphere, soil, nd vegetation to generate the thermal environment and power the hydrological and nutrient cycles (eg., energy required to evaporate water). The second category of ecological constraints limiting primary productivity involves the occurrence of rigorous abiotic factors which prevent solar energy cap- ture from being maximized. Water, temperature, and nutrient limitations frequently prevent a suificient leaf canopy from developing to intercept the available photo: Eynthetically active radiation (Lewis 1969; Begon et al. 1986). For example, plant canopies may be nonexistent for several months of the year in temperate or arid and semi-arid regions. Similarly, abiotic limitations mean that maximum photosynthetic efficiencies are seldom, if ever, attained whena plant canopy is present. Itis generally ‘estimated that less than 1% of the solar energy at the earth’s surface is converted into chemical energy by terrestrial vegetation (Lewis 1969; Begon et al. 1986). This is Gespite the fact that a conversion efficiency of approximately 20% can be realized for individual leaves under ideal environmental conditions before a biochemical limita- tion is encountered within the photosynthetic process (Lawlor 1987). Itis important tonote that the amount of solar energy captured in primary production represents the folal amount of energy available for utilization by heterotrophic organisms within the system, Secondary productivity (animal growth/area/time) is also limited by two broad categories of ecological constraints in addition to the availability of primary produc: tion The first constraint involves the inability of herbivores to consistently consume the majority of the primary production produced. Primary production varies widely through time and space, making it difficult to balance herbivore density with the fluc: tuating food resource. That portion of the herbaceous biomass available in excess of current animal demand senesces within a period of weeks following its production (Parsons et al: 1983; Chapman et al, 1984). Eventually this material enters the decom- poser compartment as litter (detrital food chain). In addition, most primary produc: tion in grasslands is located below-ground as roots and gowns making it inaccessible to large herbivores (Sims and Singh 1978b; Stanton 1988). The second category of constraints limiting secondary productivity is related toAn Ecological Perspective 15 he quality of primary production (nutritional value; see Chapter 2) ingested by he:- bivores. A substantial portion of the total energy ingested by herbivores is lost 25 methan: (in ruminants), urine, or feces, and a large portion of the metabolizable energy is utilized in basal metabolism (see Chapter 2). Itis only the remaining energy, approximately 10% of the total ingested, which is available for animal growth (Dean et al. 1975; Rode et al. 1986). Nutrient Cycling. The availability and transfer of nutrients constitutes a second indispensable function of ecological systems, Essential nutrients (carbon, nitrogen, Phosphorus, etc) form an integral component of biochemical processes and metabolic pathways within organisms which directly influence the initial capture and flow of energy through the system (Odum 1971; Wilkinson and Lowery 1973). For example, photosynthesis increases linearly over a range of leaf nitrogen concentra- tions (Field and Mooney 1986), and animal growth frequently increases with increasing nitrogen availability in the diet (Mattson 1980). However, unlike energy flow, nutrients cycle from their reservoir within the soil or atmosphere into the biotic ) component of the system (ie., producers and consumers) and then back into soil or atmospheric reservoirs within the system (Fig. 1.3) Plants initially assimilate many of the essential nutrients from the abiotic environ- ment subsequently used by animals. For example, herbivores can only acquire nitrogen by consuming plants, even though the atmosphere contains approximately 80% nitrogen by volume (Odum 1971; Wilkinson and Lowery 1973). However, as with energy, nutrients may be transferred through either the grazing or detrital food chain within the system (Fig. 1.3). Regardless of the food chain in which they are incorporated, nutrients are eventually returned to their inorganic form following organic matter decomposition by microorganisms within the decomposer compart. ment (Stanton 1988). Figure 1.3, Atmosphere Sy ow, se so A are, OTE Runt Evoren Fertilizer Dernes Soil Exsfetion Inmebitesion Mine Biecion Weatfering Fingtion : Simplified illustration of nutrient cycling within ecological systems. Nutrients move from their respective reservoirs within the abiotic component of the system, into the biotic component, and back into the environment in a cyclic pattern (after Wilkinson and Lowery 1973).GRAZING AND ECOLOGICAL SYSTEMS Plant consumption by herbivores (ie,, grazing food chain) introduces an ac tional feeding level between the primary producers and the decomposers. The objec- tive of this section is to examine how grazing influences energy flow and nutrient ng within ecological systems. A case study is presented to quantitatively illus trate the influence of grazing on energy flow and demonstrate the utility of the ecosystem concept to the investigation of grazed systems. Energy Flow ‘An Ecological Dilemma. The percentage of annual above-ground primary produc- tion utilized by herbivores varies greatly, but estimates generally range between 20% and 50% (Scott et al. 1979; Detling 1988). Although much higher levels of utilization can occur, in excess of 90%, they are generally restricted to specific regions or years. Insects and small mammals may consume as much as 10-15% of the annual above- ground production. An even smaller portion of the total annual primary production is utilized by domestic herbivores because approximately 60-90% of the production occurs below-ground in grassland systems (Sims and Singh 1978; Stanton 1988). The portion of annwal production not utilized by herbivores is eventually consumed by microorganisms in the decomposer compartment while a small portion is stored as organic compounds within a long-term carbon pool in the soil (Clark 1977). The fundamental ecological dilemma encountered in grazed systems is the inability to simultaneously optimize the interception and conversion of solar energy into primary production and the efficient harvest of primary production by her- bivores (Parsons et al. 1983). Severe grazing ensures that available production is effi- ciently harvested, but eventually reduces production by minimizing the subsequent capture of solar energy. Alternatively, lenient grazing maximizes primary produc tion, but a large percentage of the production is incorporated into the decomposer compartment without being consumed by herbivores, Contrasting patterns of energy flow between the grazing and detrital food chains are clearly illustrated in an experi- ment conducted with two perennial ryegrass pastures grazed by sheep (Fig. 1.4). One of the pastures was grazed Ieniently (24 sheep/ha) to maintain a leaf area index of approximately three (leaf area: ground area ratio of three), and the other was grazed severely (47 sheep/ha) to maintain a leaf area index of one. The total amount of energy captured by photosynthesis was 43% greater in the leniently grazed pasture as opposed to the severely grazed pasture because a greater percentage of the available Proavcer b Pesta Figure 1.4. Energy capture and flow (kg carbon/ha/day) within (a) leniently and (b) severely grazed perennial ryegrass pasture. A greater amount of solar energy is converted into Fyegrass production in the leniently grazed pasture, but this grazing regime reduces » the relative amount of energy consumed by livestock and incresses the relative amount of energy transferred into the decomposer compartment in comparison with the severely grazed pasture (after Parsons et al. 1983). FoagrseeAn Ecological Perspective 17 solar radiation was intercepted by the plant canopy. However, animal consumption was 40% greater in the severely grazed pasture than in the leniently grazed pasture, even though production was less, because of greater livestock numbers per hectare. The end result was that 42% of the energy captured by ryegrass in the leniently grazed pasture entered the detrital food chain, while only 13% was consumed by livestock By contrast, only 26% of the energy captured by ryegrass in the severely grazed pasture entered the detrital food chain, while 25% entered the grazing food chain. The remainder of the energy captured by photosynthesis (49%) was either allocated to root growth or used in plant respiration. These data illustrate that primary production and efficient biomass utilization cannot be maximized simultaneously because of the contribution of leaf area to both processes A Case Study. The influence of grazing on energy flow within ecological systems can best be illustrated quantitatively by evaluating a simple case study from the Texas Experimental Ranch near Throckmorton, Texas. This mixed-grass prairie site was grazed continuously throughout the year at a stocking rate considered severe for the region. The system yielded a mean above-ground herbaceous production of 3183 ) kg/ha/yr (Heitschmidt et al. 19872), individual animal gains of 200 kg/cow, and livestock gains of 53.5/kg/ha/yr (Heitschmidt et al. 1990). These production values can easily be converted into energy values by virtue of the fact that 1 kg of plant and animal tissue contains approximately 19.7 and 23.5 MJ (mega [million] joules) respectively (Golley 1961; Odum 1971). (A joule [J] is.the expression for energy in the International System of Units; 1 calorie = 4.19 J; 1] = 0.24 calories.) The total amount of solar energy received annually at the site provides a convenient reference point with which to compare energy values at various loca~ tions within the system. The total input of solar energy at the latitude of the Experi- mental Ranch (33° 20'N) is.approximately 63,000,000 MJ/ha/yr (Cinguemani et al. 1978). In this system, approximately 62,705 MJ/ha/yr is captured in herbaceous above-ground vegetation; 313,525 MJ/ha/yr is captured in total (above- and below- ground) herbaceous vegetation (assuming 80% of the primary production is below- ground; Stanton 1988); and 1,257 Mj/ha/yr is transferred into livestock gains (Fig. 15). Decreasing energy values from the initial input of solar énergy to vegetation and finally to livestock gains clearly demonstrate the inefficiency of energy transfer within ) the system. System Component Energy Conversion Efficiency Content Solr Enerty Tere, 62,000.000 4 Photosynthetic Rediation 28,000,000 nuiow nr Pein fous : : oso% ‘rovegound 62.708 002% Betongrund 2s0820 20% Tera! 313.525 Seeendory reduction esr Figure 1.5. Energy content (MJ/ha/ys) and transfer efficiencies (2%) for primary and secondary production in relation to total and photosynthetically active solar radiation at the ‘Texas Experimental Ranch (from Heitschmidt et al. 1987a; 1990). Energy values are calculated by multiplying primary and secondary production values by 19.7 and 23.5 MJ/kg, respectively. Conversion efficiencies represent the quotient of two energy values at specified locations within the system multiplied by 100.18 D.D. Briske and R. K. Heitschmidt An important aspect of energy flow analysis is that transfer efficiencies can be culated by dividing the amount of energy captured within one level of the system by mount of energy in a preceding level (Golley 1960; Pimm 1988). Above- ground herbaceous vegetation captured 0.10% and 0.22% of the total solar radiation and photosynthetically active radiation/ha/yr respectively (Fig. 1.5). A slightly greater conversion efficiency, 0.50%, is observed when total (above- and below- ground) herbaceous annual production is considered. Conversion efficiencies decrease even further when energy transfer into livestock production is calculated pecause of the energy loss which occurs between feeding levels. Approximately 0.002% of the energy available in total annual solar radiation is transferred into livestock gains in comparison with 2% of the energy available in above-ground production. These minimal conversion efficiencies, which progressively decrease with the incorporation of additional feeding levels, are similar to those estimated in other grassland systems (Macfadyen 1964; Coleman et al. 1976; Snayden 1981; Akiyama et al. 1984). ‘The intrinsic inefficiencies of energy flow in grazed systems should not be inter- preted to imply that these systems possess an insignificant potential for secondary production. Grazing could potentially yield an estimated 7.5 trillion MJ of animal production annually from grasslands and savannas of the world, assuming that large herbivores consume 20% of the above-ground production and possess a conversion efficiency of 2% (Fig. 1.5). These estimates demonstrate the tremendous importance of grazing to the human food supply on a global basis. Substantial increases in secondary production can be attained with only modest increases in ecological effi- Giencies resulting from effective management strategies. A combined increase of only 0.01% in harvest and conversion efficiencies would potentially increase secondary production by 75 billion MJ. Nutrient Cycling Grazing may also modify the rate and pattern of energy flow in ecological systems by influencing nutrient availability. Nutrient availability, in turn, governs the efficiency with which organisms acquire and process energy. Grazing affects nutrient cycling by accelerating the rate of nutrient conversion from an organic form (amino acids and proteins) to an inorganic form (nitrate and ammonium), This process, termed mineralization, is critical to grassland production because a large proportion of the essential nutrients are bound in organic matter within the soil (Wilkinson and Lowrey 1973; Woodmansee et al. 1978). However, only those nutrients in specific inorganic forms are available for plant absorption. Grazing increases mineralization by reducing the particle size of plant material (eg, chewing and rumination) and providing a favorable environment for microbial activity (eg, high body temperatures; Wilkinsori and Lowrey 1973; Floate 1981). Yet herbivores retain only a small portion of the nutrients consumed, thereby rapidly returning most nutrients to the system in urine and feces. Nutrients excreted in urine—primerily nitrogen, potassium, magnesium, and sulfur—are in the inorganic form and therefore immediately available for plant absorption (Wilkinson and Lowsey 1973). In contrast, a greater proportion of nutrients in fecal material and ungrazed plant material than in urine are bound in organic compounds and must be mineralized by decomposers prior to plantabsorption. Consequently, a portion of the nutrients incorporated into primary production become available for reabsorption more rapidly when transferred through the grazing food chain than when trans- ferred directly into the decomposer compartment (Wilkinson and Lowrey 1973;An Ecological Perspective 19 Floate 1981). Estimates of higher nutrient concentrations in vegetation of grazed than of ungraved systems support the assumption of increased rates of nutrient cycling (Detling 1988). Nutrient transfer through the grazing food chain potentially increases the rate of cycling, but in so doing, it also increases the potential for nutrient losses from the system (Woodmansee 1978; Floate 1981). Nutrient losses from grazed systems primarily occur as volatilization, leaching, soil erosion, and livestock removal from the system (Wilkinson and Lowrey 1973; Woodmansee et al. 1981). Nutrient losses are highly variable and are influenced by a large number of environmental variables inchiding nutrient solubility, soil morphology and chemistry, climate, and topography (see Chapter 5). Nutrient losses associated with herbivore removal from the system are minimized by the limited productivity of many grasslands and the digestive physiology of herbivores (Floate 1981). Nutrient availability i