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With Clinical Disease : Fescue Foot and Perennial Ryegrass Staggers

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With Clinical Disease : Fescue Foot and Perennial Ryegrass Staggers AN ABSTRACT OF THE THESIS OF JOHN TOR-AGBIDYE for the degree of Master of Science in Veterinary Science presented on August 13, 1993. Title:Correlation of Endophyte Toxins (Ergovaline and Lolitrem B) with Clinical Disease: Fescue Foot and Perennial Ryegrass Staggers Abstract Approved:Redacted for Privacy Dr. A. Morrie Craig Endophytic fungi (A. coenophialum and A. lolii) which infect grasses produce ergot alkaloids that serve as the grasses' chemical defenses and enhance the vigor of the grass.Turf-type tall fescue with high endophyte levels has been deliberately developed to produce a greener, more vigorous, pest-resistant turf. Consumption of endophyte-infected grass causes various toxicity symptoms in livestock. Cattle in the southeastern and midwestern United States, where tall fescue is grown on 14 million hectares, often develop signs of toxicosis during summer months from grazing plants in fected by A. coenophialum. A more severe form of the disease, fescue foot, has been associated with cold environment and reported in late fall and winter months not only in the southeastern United States but also in the northwest United States.In New Zealand, where perennial ryegrass is grown on 7 million hectares of pasture, sheep often develop a condition called ryegrass staggers from grazing plants infected by A. lolii. New Zealand reports economic losses grazing plants infected by A. lolii. New Zealandreports economic losses associated with the sheep industry of $205 millionper year.In the United States, economic losses associated with the beef cattle industry alone is estimatedat $600 million per year. Range finding experiments and case studies of fescue foot and perennial ryegrass staggers (PRGS) were conducted on cattle and sheep under grazing and barn conditions. The main objective was to determine threshold levels ofthe endophyte toxins, ergovaline (EV) (appendix 1) and lolitrem B (appendix 2),associated with the diseases of fescue foot and PRGS respectively. Fescue footwas experimentally induced in cattle under barn studies in the spring with 825 ppb ergovaline.The ergovaline contaminated feed was given fora period of 42 days. Similar barn studies in sheep in spring to early summer did not produce clinicalfescue foot with up to 1215 ppb. Field studies of natural fescue foot in a herd of sheepwere conducted, (ie 540 ppb) values of ergovaline in the feed, but clinical diseasewas not produced in late fall through winter. A case study froma herd of sheep revealed 813 ppb dietary ergovaline had produced fescue foot in the months of fall(November). Fields of perennial ryegrass (PRG) where sheep received 2,135 ppb lolitremB toxin were associated with clinical cases of PRGS in 42 sheep of 237sheep (18 percent incidence rate) in the Willamette Valley of Oregon. Three months later, sheep on this same field which then had 1,465 ppb lolitrem B, didnot have PRGS. These were the first range finding experiments undertaken in thislocale to document threshold levels of endophyte toxins associated with fescue foot andPRGS. Correlation of Endophyte Toxins (Ergovaline and Lolitrem B) with Clinical Disease: Fescue Foot and Perennial Ryegrass Staggers by John Tor-Agbidye, DVM A THESIS submitted to Oregon State University In partial fulfillment of the requirements for the degree of Master of Science Completed August 13, 1993 Commencement June 1994 APPROVED: Redacted for Privacy Profes eterinary Medicine in char of Major Redacted for Privacy Redacted for Privacy Dean of Gradu Date thesis is presented August 13, 1993 Typed by John Tor-Agbidye TABLE OF CONTENTS INTRODUCTION 1 LITERATURE REVIEW 7 Tall Fescue 8 Endophyte Location in PlantLevels of Toxin 9 Biosynthesis and Mechanism of Distribution of Ergovaline, Lolitrem B and Related Metabolites (Peramine) in Endophyte-infected (E+) Plants 11 Etiology of Fescue Toxicosis 12 Nature and Mode of Action of Toxins 12 A Review of the Hormone Prolactin and Neurotransmitters (dopamine and serotonin) 14 Dopamine as a Neurotransmitter 17 Glutamate as a Neurotransmitter 18 Serotonin as a Neurotransmitter 18 Disease Conditions Produced 19 Fescue Foot 20 Fat Necrosis 20 Agalactia 21 Summer Fescue Toxicosis 21 Fescue foot 22 The Pasture in Relation to Fescue Foot 22 Signs of Fescue Foot 23 Blood Profiles 27 Blood Flow and Thermoregulation 28 Diseases with Signs Similar to Fescue Foot 29 Treatment and Prevention of Fescue Foot 30 Management Cure 30 Food from Affected Animals 31 Research on Naturally Occurring Cases of Fescue Foot 31 Effects of Fescue Toxicosis on Reproduction in Livestock 36 Puberty 40 Milk Production 41 Horses 41 Fertility 43 Mechanisms of Toxicity 45 Implications 46 Summary of Research on Tall Fescue 47 Tall Fescue Problem Research- Major Research Objective 1 47 Tall Fescue Problem Research- Major Research Objective 2 48 Tall Fescue Problem Research- Major Research Objective 3 48 Tall Fescue Problem Research-- Major Research Objective 4 48 Management 49 New and Improved Varieties 50 The Future 50 Tall Fescue Toxicosis: Could there be a public health problem here? 50 Perennial Ryegrass 53 Areas Reporting Perennial Ryegrass Staggers 53 Etiology of Perennial Ryegrass Staggers 54 Nature and Mode of Action of Toxins 54 Signs of Perennial Ryegrass Staggers 57 Disease Conditions Similar to Perennial Ryegrass Staggers 58 Management 59 MATERIALS AND METHODS 62 Experiment 1:Field Studies with Sheep on E+ Tall Fescue Pastures 62 Pasture 1 62 Pasture 2 63 Experiment 2: Barn Studies 64 Barn Study on EV Toxin and Sheep 64 Barn Study on EV Toxin and Cattle 65 Case Report 66 Field Outbreak of Fescue Foot in Sheep 66 Field Outbreak of Perennial Ryegrass Staggers in Sheep 67 Protocol for Endophyte Analysis 68 Protocol for Ergovaline Analysis 68 Reagents 69 Solutions 69 Instrumentation 69 Mobile phase 70 Sample collection 70 Sample preparation 70 Standards 72 Protocol for Prolactin Analysis 72 Protocol for Lolitrem B Analysis 73 Mobile Phase 73 Gradient program 74 Solutions 74 Hematology 75 Blood Chemistry 75 Statistical Analysis 75 RESULTS 77 Experiment 1:Field Studies with Sheepon E+ Tall Fescue Pastures 77 Pasture 1 77 Pasture 2 78 Experiment 2: Barn Studies 79 Barn Studies on Sheep 79 Barn Studies on Cattle 80 Case Report 81 Field Outbreak of Fescue Foot in Sheep 81 Field Outbreak of Perennial Ryegrass Staggers in Sheep 82 DISCUSSION 91 Tall Fescue Studies 91 Perennial Ryegrass Studies 97 Conclusions 104 BIBLIOGRAPHY 106 APPENDIX I 119 II 120 Glossary 121 LIST OF FIGURES Figure Page 1. Serum prolactin of ewes grazed on E+ field 1 at 350 ppb EV 85 2. Serum prolactin of ewes grazed on E+ field 2 86 at 500 ppb EV 3. Ergovaline distribution studies on dormant 87 tall fescue 4. Studies with ewes on endophyte-infected 88 tall fescue field 5. Barn studies with sheep on increasing dietary 89 EV levels 6. a) One of the cattle showed signs of fescue foot 90 b) Cornification and elongation of hooves due to vasoconstrictive impairment of blood flow to the hoof area Appendices I Ergovaline 119 II Lolitrem B 120 LIST OF TABLES Table Page 1. Hematologic values in experimental sheep fed 1,215 ppb dietary ergovaline 83 2. Serum biochemical values of experimental 84 sheep fed 1,215 ppb dietary ergovaline Correlation of Endophyte Toxins (Ergovaline and Lolitrem B) with Clinical Disease: Fescue Foot and Perennial Ryegrass Staggers INTRODUCTION Many species of grasses (Gramineae) of the subfamily Poodeae (Festucoidae) are associated with intercellular fungi. The fungi grow endophytically within seeds, leaves, culms, rhizomes, and meristems of grasses and never show external signs of infection nor signs of a disease. The fungus associated with thesegrasses belongs to the anamorphic genus Acremonium link section and consists primarily of two species: A. coenophialum and A. lolii. These two fungal species are perennially associated with tall fescue (Festuca arundinacea Schreb.) or perennialryegrass (Loilium perenne L.), and current evidence indicates that these symbiota are mutualisms. Unlike other symbiota, once the association is established, the fungal andgrass components do not live apart, even briefly; therefore, these associations are describedas an obligately conjunctive symbiota. Fungal mutualism of the symbiota produce toxic secondary metabolites which affect a broad range of invertebrate and vertebrate animals, as wellas microorganisms. Fungal mutualists are also associated with mechanisms which allow the symbiota to tolerate many severe or limiting abiotic environmental conditions. These symbiota apparently established their associations early during evolution, but did not co-evolve with their hosts. Much of their competitive ability isa direct outcome of the many secondary products produced by the fungus. Endophytic fungi (A. coenophialum) which infect tall fescue produce chemical 2 (ergot) alkaloids that serve as the grass's chemical defenses and enhance the vigor of the grass. The principal animal toxin in tall fescue is believed to be ergovaline (EV) (see Appendix 1).Turf-type tall fescue with high endophyte levels has been deliberately developed to produce a greener,more vigorous, pest-resistant turf. The ergot alkaloids not only inhibit plant pests and diseases but also are toxic to large animals. Consumption of endophyte-infected tall fescuecauses various toxicity signs in livestock. These include summer fescue toxicosis (poor growth, increased sensitivity to heat stress), fescue foot, foot necrosis, and impaired reproduction. These effects are largely (or perhaps totally) due to alterations in secretion ofthe
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