Browsing Journal of Range Management, Volume 54, Number 4 (July 2001) by Subjects
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Alkaloids as anti-quality factors in plants on western U.S. rangelandsAlkaloids constitute the largest class of plant secondary compounds, occurring in 20 to 30% of perennial herbaceous species in North America. Alkaloid-containing plants are of interest, first because alkaloids often have pronounced physiological reactions when ingested by livestock, and second because alkaloids have distinctive taste characteristics. Thus, alkaloids may kill, injure, or reduce productivity of livestock, and have the potential to directly or indirectly alter diet selection. We review 7 major categories of toxic alkaloids, including pyrrolizidine (e.g., Senecio), quinolizidine (e.g., Lupinus), indolizidine (e.g., Astragalus), diterpenoid (e.g., Delphinium), piperidine (e.g., Conium), pyridine (e.g., Nicotiana), and steroidal (Veratrum-type) alkaloids. Clinically, effects on animal production vary from minimal feed refusal to abortion, birth defects, wasting diseases, agalactia, and death. There are marked species differences in reactions to alkaloids. This has been attributed to rumen metabolism, alkaloid absorption, metabolism, excretion or directly related to their affinity to target tissues such as binding at receptor sites. In spite of alkaloids reputed bitter taste to livestock, some alkaloid-containing plant genera (e.g., Delphinium, Veratrum, Astragalus, Oxytropis, and Lupinus) are often readily ingested by livestock. Management schemes to prevent losses are usually based on recognizing the particular toxic plant, knowing the mechanism of toxicity, and understanding the temporal dynamics of plant alkaloid concentration and consumption by livestock. Once these aforementioned aspects are understood, losses may be reduced by maintaining optimal forage conditions, adjusting grazing pressure and timing of grazing, aversive conditioning, strategic supplementation, changing livestock species, and herbicidal control.
Anti-quality components in forage: Overview, significance, and economic impactAlthough recognized in importance from the dawn of history, forages have too often been underestimated and undervalued perhaps in part because animal performance has frequently failed to reflect apparent forage quality. Anti-quality components, diverse impediments to quality, have evolved as structural components and as secondary metabolites. They include mineral imbalances or can be related to the presence of insects and diseases. Animal behavior and adaptation are increasingly recognized as important aspects of anti-quality factors. An anti-quality component may reduce dry matter intake, dry matter digestibility, or result in nutritional imbalances in animals. They can act as a direct poison compromising vital systems, result in abnormal reproduction, endocrine function, and genetic aberrations, trigger undesirable behavior responses, or suppress immune function leading to increased morbidity and mortality. The economic impact of anti-quality factors on individual herds can be devastating but definable. Broadscale economic impacts of anti-quality factors are far more difficult to estimate. A loss of 0.22 kg/day in potential gain of stocker cattle due to anti-quality factors during a 166-day grazing season translates into a loss of about 55/steer at 1.45/kg or over 2 billion annually when applied to the U.S stocker cattle. Economic losses to tall fescue (Festuca arundinacea Schreb.) toxicosis in the U.S. beef industry are probably underestimated at 600 million annually. Reproductive and death losses of livestock due to poisonous plants have been estimated at 340 million in the 17 western states alone. These examples of economic losses due to anti-quality factors may be upper bounds of actual losses but even if a small proportion of the expected losses were eliminated through research, the potential payoff would be extremely high.
Herbivore response to anti-quality factors in foragesPlants possess a wide variety of compounds and growth forms that are termed "anti-quality" factors because they reduce forage value and deter grazing. Anti-quality attributes can reduce a plant's digestible nutrients and energy or yield toxic effects. Herbivores possess several adaptive mechanisms to lessen the impacts of anti-quality factors. First, herbivores graze selectively to limit consumption of potentially harmful plant compounds. Grazing animals rely on a sophisticated system to detect plant nutritional value or toxicity by relating the flavor of a plant to its positive or negative digestive consequences. Diet selection skills are enhanced by adaptive intake patterns that limit the deleterious effects of plant allelochemicals; these include cautious sampling of sample new foods, consuming a varied diet, and eating plants in a cyclic, intermittent, or carefully regulated fashion. Second, grazing animals possess internal systems that detoxify or tolerate ingested phytotoxins. Animals may eject toxic plant material quickly after ingestion, secrete substances in the mouth or gut to render allelochemicals inert, rely on rumen microbes to detoxify allelochemicals, absorb phytochemicals from the gut and detoxified them in body tissues, or develop a tolerance to the toxic effects of plant allelochemicals. Understanding the behavioral and metabolic abilities of herbivores suggests several livestock management practices to help animals contend with plant anti-quality characteristics. These practices include offering animals proper early life experiences, selecting the appropriate livestock species and individuals, breeding animals with desired attributes, and offering nutritional or pharmaceutical products to aid in digestion and detoxification.
Lignin and fiber digestionLignin is a polymer formed from monolignols derived from the phenylpropanoid pathway in vascular plants. It is deposited in the cell walls of plants as part of the process of cell maturation. Lignin is considered an anti-quality component in forages because of its negative impact on the nutritional availability of plant fiber. Lignin interferes with the digestion of cell-wall polysaccharides by acting as a physical barrier to microbial enzymes. Lignification therefore has a direct and often important impact on the digestible energy (DE) value of the forage. There are a number of plant-related factors that affect lignification in individual plants and plant communities. Lignification is under genetic control and there are considerable differences in lignin concentration and composition among species and even genotypes within species. Genetic differences in lignification are first expressed at the cellular level and are affected by biochemical and physiological activities of the cell. As cells differentiate, differences in lignification occur depending on the tissues and organs being developed. Lignification tends to be most intense in structural tissues such as xylem and sclerenchyma. Plant organs containing high concentrations of these tissues, such as stems, are less digestible than those containing lower concentrations. The relative proportion of lignified tissues and organs typically increases as plants mature so there is often a negative relationship between digestibility and maturity. All of these plant processes respond to environmental factors that can affect the extent and impact of lignification. Temperature, soil moisture, light, and soil fertility can have either direct or indirect effects on lignification. The most useful management practices for minimizing the negative effects of lignification are manipulation of the plant community such that it contains more desirable species and harvest management to maintain plants in a vegetative stage of development.