Forages and grazing in horse nutrition pp 17-43 | Cite as
Grass physiology and its relation to nutritive value in feeding horses
Abstract
Forage is the primary feed of the horse; it normally comprises more than 50% of the horse’s diet on DM basis and it may supply 100% of the ration of many horse categories. Grasses are well adapted to frequent defoliation and to the presence of large herbivores, and consequently they cover, globally, large areas of natural and seminatural vegetation, and are also widely used in intensive forage production. As the Graminae is one of the largest plant families — over 600 genera — there is a large variation in the physiological mechanisms by which grasses react to environmental variables and management factors. In addition, humans utilize grass yield in several ways, i.e. as grazed grass, silage, haylage or hay. Consequently, crop physiology behind the grass yield is vast. In this paper we first describe on outline of plant physiology of grasses and main differences between C3 and C4 grasses as well as difference between forage legumes and grasses. Secondly, we focus on plant physiology of grasses that is of most relevance in the context of equine nutrition. We present an outline of key processes affecting the digestibility of grasses, since digestibility is the most important single feature of forage affecting the nutritive value. In addition, we cover fructan metabolism in grasses and its consequences in producing pasture, silage, haylage and hay. We also present a summary of effects of nitrogen on the production and nutritive value of grasses. We conclude that knowledge of plant physiology provides tools to understand the changes in forage quality and quantity. Based on this knowledge we have tools to choose the most proper management options in order to produce high quality forage for different types of horses.
Keywords
crop physiology equine nutrition nutritive value digestibility graminae fructan carbohydrate nitrogenNotes
Acknowledgements
The authors wish to thank Maarit Hyrkas and Alan Hopkins for their excellent help in preparing the manuscript.
References
- Aiken, G.E., D.I. Bransby and C.A. McCall, 1993. Growth of yearling horses compared to steers on high-endophyte and low-endophyte infected tall fescue. J. Equine Vet. Sci. 13, 26–28.CrossRefGoogle Scholar
- Akin, D.E., 1989, Histological and physical factors affecting digestibility of forages. Agron. J. 81, 17–25.CrossRefGoogle Scholar
- Akin, D.E., S.L. Fales, L.L. Rigsby and M.E. Snook, 1987. Temperature effects on leaf anatomy, phenolic-acids, and tissue digestibility in tall fescue. Agron. J. 79, 271–275.CrossRefGoogle Scholar
- Bélanger, G. and R.E. McQueen, 1997. Leaf and stem nutritive value of timothy cultivars differing in maturity. Can. J. Plant Sci. 77, 237–245.CrossRefGoogle Scholar
- Bélanger, G., 1998. Morphogenetic characteristics of timothy grown with varying N nutrition. Can. J. Plant Sci. 78, 103–108.CrossRefGoogle Scholar
- Bélanger, G., G.F. Tremblay and D. Mongrain, 2008. Yield and nutritive value of the spring growth of an ageing timothy sward. Can. J. Plant Sci. 88, 457–464.CrossRefGoogle Scholar
- Bertrand, A., G.F. Tremblay, S. Pelletier, Y. Castonguay and G. Bélanger, 2008. Yield and nutritive value of timothy as affected by temperature, photoperiod and time of harvest. Grass Forage Sci. 63, 421–432.CrossRefGoogle Scholar
- Blevins, D.G. and D.J. Barker, 2007. Nutrients and water in forage plants. In: Barnes, R.F., C.J. Nelson, K.J. Moore and M. Collins (eds.) Forages: The science of grassland agriculture. 6th ed. Blackwell Publishing, Oxford, UK, p. 67–80.Google Scholar
- Bonesmo, H. and G. Bélanger, 2002. Timothy yield and nutritive value by the CATIMO model: I. Growth and nitrogen. Agron. J. 94, 337–345.Google Scholar
- Booysen, P.D. and C.J. Nelson, 1975. Leaf area and carbohydrate reserves in regrowth of tall fescue. Crop Sci. 15, 262–266.CrossRefGoogle Scholar
- Briske, D.D., 1985. Plant response to defoliation: Morphological considerations and allocation priorities. In: Joss, P.J., P.W. Lynch and O.B. Williams (eds.) Rangelands, a resource under siege. Cambridge University Press, Cambridge, UK, p. 425–427.Google Scholar
- Brito, A.F., G.F. Tremblay, A. Bertrand, Y. Castonguay, G. Bélanger, R. Michaud, H. Lapierre, C. Benchaar, H.V. Petit, D.R. Ouellet and R. Berthiaume, 2008. Alfalfa cut at sundown and harvested as baleage improves milk yield of late-lactation dairy cows. J. Dairy Sci. 91, 3968–3982.PubMedCrossRefGoogle Scholar
- Bush, L., C.A. Roberts and C. Schultz, 2007. Plant chemistry and antiquality components in forage. In: Barnes, R.F., C.J. Nelson, K.J. Moore and M. Collins (eds.) Forages: the science of grassland agriculture. 6th ed. Blackwell Publishing, Oxford, UK, p. 509–528.Google Scholar
- Buxton, D.R. and S.L. Fales, 1994. Plant environment and quality. p. 155–199. In: G.C. Fahey (eds.) Forage quality, evaluation and utilization. ASA-CSSA-SSSA, Madison, WI.Google Scholar
- Calviere, I. and M. Duru, 1995. Leaf appearance and senescence patterns of some pasture species. Grass Forage Sci. 50, 447–451.CrossRefGoogle Scholar
- Casler, M.D. and R.L. Kallenbach, 2007. Cool-season grasses for humid areas. In: Barnes, R.F., C.J. Nelson, K.J. Moore and M. Collins (eds.) Forages: the science of grassland agriculture. 6th ed. Blackwell Publishing, Ames, Iowa, USA, p. 211–220.Google Scholar
- Chatterton, N.J., P.A. Harrison, J.H. Bennet, and K.H. Asay, 1989. Carbohydrate portioning in 185 accessions of graminae grown under warm and cool temperature. J. Plant Physiol. 134, 169–179.CrossRefGoogle Scholar
- Chatterton, N.J., P.A. Harrison, W.R. Thornley and E.A. Draper, 1990. Oligosaccharides in foliage of agropyron, bromus, dactylis, festuca, lolium and phleum. New Phytol. 114, 167–171.CrossRefGoogle Scholar
- Chen, L., C.K. Auh, F. Chen, X.F. Cheng, H. Aljoe, R.A. Dixon and Z.Y. Wang, 2002. Lignin deposition and associated changes in anatomy, enzyme activity, gene expression, and ruminal degradability in stems of tall fescue at different developmental stages. J. Agric. Food Chem. 50, 5558–5565.PubMedCrossRefGoogle Scholar
- Ciavarella, T., R. Simpson, H. Dove, B. Leury and I. Sims, 2000. Diurnal changes in the concentration of water-soluble carbohydrates in Phalaris aquatica L. pasture in spring, and the effect of short-term shading. Aust. J. Agric. Res. 51, 749–756.CrossRefGoogle Scholar
- Coenen, M., I. Vervuert and A. Moessler, 2005. Hydrogen and methan exhalation profiles of horses after consumption of grass meal (cellulose), sugar beet pulp (pectin), oats (starch) and jerusalem artichike (fructan). Proc. 19th Symp. Equine Sci. Soc.158-159.Google Scholar
- Collins, M. and W.K. Coblentz, 2007. Post-harvest physiology. In: Barnes, R.F., C.J. Nelson, K.J. Moore and M. Collins (eds.) Forages: the science of grassland agriculture. 6th ed. Blackwell Publishing, Oxford, UK, p. 583–599.Google Scholar
- Cruz, P. and M. Boval, 2000. Effect of nitrogen on some morphogenetic traits of temperate and tropical perennial forage grasses. In: Lemaire, G., J. Hodgson, A. de Moraes, P.C. de F. Carvalho and C. Nabinger (eds.), Grassland ecophysiology and grazing ecology. CABI Publishing, Wallingford, UK, p. 151–168.CrossRefGoogle Scholar
- Cudderford, D., 2004. Voluntary food intake by horses. In: Juliand, V. and W. Martin-Rosset (eds.) Nutrition of the performance horse: which system in Europe for evaluating the nutritional requirements? EAAP Publication No. 111. Wageningen Academic Publishers, Wageningen, the Netherlands, pp. 89–100.Google Scholar
- Cymbaluk, N.F., 1990. Comparison of forage digestion by cattle and horses. Can. J. Anim. Sci. 70, 601–610.CrossRefGoogle Scholar
- Davies, A., 1988. The regrowth of grass swards. In: Jones, M.B. and A. Lazenby (eds.) The grass crop. Chapman and Hall Ltd, London, UK, p. 85–127.CrossRefGoogle Scholar
- Deinum, B., J. Debeyer, P.H. Nordfeldt, A. Kornher, O. Ostgard and G. Vanbogaert, 1981. Quality of herbage at different latitudes. Neth. J. Agric. Sci. 29, 141–150.Google Scholar
- Donaghy, D. and W. Fulkerson, 1997. The importance of water-soluble carbohydrate reserves on regrowth and root growth of Lolium perenne (L.). Grass Forage Sci. 52, 401–407.CrossRefGoogle Scholar
- Doreau, M., C.Moretti and W. Martin-Rosset, 1990. Effect of quality of hay given to mares around foaling on their voluntary intake and foal growth. Ann. Zootech., 39, 125–131.CrossRefGoogle Scholar
- Duru, M. and I. Calviere, 1996. Influence of leaf area index and carbohydrate reserves at cutting on the leaf growth of two forage grasses (Dactylis glomerata L and Festuca arundinaceae schreb). Can. J. Plant Sci. 76, 269–276.CrossRefGoogle Scholar
- Duru, M., 1997. Leaf and stem in vitro digestibility for grasses and dicotyledons of meadow plant communities in spring. J. Sci. Food Agric. 74, 175–185.CrossRefGoogle Scholar
- Earle, W.E., D.L. Cross, L.W. Hudson, L.M. Redmond and S.W. Kennedy, 1990. Effect of energy supplementation on gravid mares grazing endophyte-infected fescue. J. Equine Vet. Sci. 10, 126–130.CrossRefGoogle Scholar
- Eduard, N., G. Fleurance, W. Martin-Rosset, P. Duncan, J.P. Dulphy, S. Grange, R. Baumont, H. Dubroeucq, F.J. Perez-Barberia and I.J. Gordon, 2008. Voluntary intake and digestibility in horses: effect of forage quality with emphasis on individual variability. Animal 2, 1526–1533.Google Scholar
- Edwards, G.R., A.J. Parsons, S. Rasmussen, and R.H. Bryant, 2007. High sugar ryegrasses for livestock systems in New Zealand. Proceedings of the New Zealand Grassland Association 69, 161–171.Google Scholar
- Fales, S.L. and J.O. Fritz, 2007. Factors affecting forage quality. In: Barnes, R.F., C.J. Nelson, K.J. Moore and M. Collins (eds.) Forages: the science of grassland agriculture. 6th ed. Blackwell Publishing, Oxford, UK, p. 569–580.Google Scholar
- Fisher, D., H. Mayland and J. Burns, 1999. Variation in ruminants’ preference for tall fescue hays cut either at sundown or at sunup. J. Anim. Sci. 77, 762–768.PubMedGoogle Scholar
- Fisher, D., J. Burns and H. Mayland, 2005. Ruminant selection among switchgrass hays cut at either sundown or sunup. Crop Sci. 45, 1394–1402.CrossRefGoogle Scholar
- Fulkerson, W.J. and K. Slack, 1995. Leaf number as a criterion for determining defoliation time for Lolium-perenne. 2. Effect of defoliation frequency and height. Grass Forage Sci. 50, 16–20.CrossRefGoogle Scholar
- Gastal, F. and C.J. Nelson, 1994. Nitrogen use within the growing leaf blade of tall fescue. Plant Physiol. 105, 191–197.PubMedGoogle Scholar
- Gastal, F. and J-L. Durand, 2000. Effects of nitrogen and water supply on N and C fluxes and partitioning in defoliated swards. In: Lemaire, G., J. Hodgson, A. de Moraes, P.C. de F. Carvalho and C. Nabinger (eds.) Grassland ecophysiology and grazing ecology. CABI Publishing, Wallingford, UK, p. 15–39.CrossRefGoogle Scholar
- Geor, R.J., 2008. Metabolic predispositions to laminitis in horses and ponies: Obesity, insulin resistance and metabolic syndromes. J. Equine Vet. Sci. 28, 753–759.CrossRefGoogle Scholar
- Gonzalez, B., J. Boucaud, J. Salette, J. Langlois and M. Duyme, 1989. Changes in stubble carbohydrate content during regrowth of defoliated perennial ryegrass (Lolium-perenne L) on 2 nitrogen levels. Grass Forage Sci. 44, 411–415.CrossRefGoogle Scholar
- Gordon, M.E., M.L. Jerina, L.K. Scott, K.E. Davison, J.K. Young and R.H. Raub, 2007. The effects of non-structural carbohydrate content and feeding rate on glucose and insulin response to meal feeding in equine. J. Equine Vet. Sci. 27, 489–493.CrossRefGoogle Scholar
- Grabber, J.H., J. Ralph, C. Lapierre and Y. Barriere, 2004. Genetic and molecular basis of grass cell-wall degradability. I. Lignin-cell wall matrix interactions. C. R. Biologies 327, 455–465.Google Scholar
- Grässler, J. and U. Borstel, 2005. Fructan content in pasture grasses. Proceedings of Equine Nutrition Conference, Hannover, Germany, p.75-76.Google Scholar
- Greenwood, D.J., G. Lemaire, G. Gosse, P. Cruz, A. Draycott and J.J. Neeteson, 1990. Decline in percentage N of C3 and C4 crops with increasing plant mass. Ann. Botany 66, 425–436.Google Scholar
- Grotelueschen, R.D. and D. Smith, 1967. Determination and identification of nonstructural carbohydrates removed from grass and legume tissue by various sulfuric acid concentrations, takadiastase, and water. J. Agr. Food. Chem. 15, 1048–1051.CrossRefGoogle Scholar
- Guay, K.A., H.A. Brady, V.G. Allen, K.R. Pond, D.B. Wester, L.A. Janecka and N.L. Heninger, 2002. Matua bromegrass hay for mares in gestation and lactation. J. Anim. Sci., 80, 2960–2966.PubMedGoogle Scholar
- Hanna, W.W. and L.E. Sollenberger, 2007. Tropical and subtropical grasses. In: Barnes, R.F., C.J. Nelson, K.J. Moore and M. Collins (eds.), Forages: The science of grassland agriculture. 6th ed. Blackwell Publishing, Oxford, UK, p. 245–255.Google Scholar
- Harris, P., S. Bailey, J. Elliott and A. Longland, 2006. Countermeasures for pasture-associated laminitis in ponies and horses. J. Nutr. 136, 2114S-2121S.PubMedGoogle Scholar
- Harris, P.A., 2005. Influence of feed and feeding on incidence of laminitis. Pfredeheilkunde 21, 64–65.Google Scholar
- Hatfield, R.D., H-J.G. Jung, G. Broderick and T.C. Jenkins, 2007. Nutritional chemistry of forages. In: Barnes, R.F., C.J. Nelson, K.J. Moore and M. Collins (eds.) Forages: the science of grassland agriculture. 6th ed. Blackwell Publishing, Oxford, UK, p. 467–485.Google Scholar
- Hay, R.K.M. and J.R. Porter, 2006. The physiology of crop yield. Blackwell Publishing, Oxford, UK.Google Scholar
- Heide, O.M., 1994. Control of flowering and reproduction in temperate grasses. New Phytol. 128, 347–362.CrossRefGoogle Scholar
- Heide, O.M., R.K.M. Hay and H. Baugerod, 1985. Specific daylength effects on leaf growth and dry-matter production in high-latitude grasses. Ann. Botany 55, 579–586.Google Scholar
- Hodgson, J. and S.C. Da Silva, 2000. Sustainability of grazing systems: Goals, concepts and methods. In: Lemaire, G., J. Hodgson, A. de Moraes, P.C. de F. Carvalho and C. Nabinger (eds.) Grassland ecophysiology and grazing ecology. CABI Publishing, Wallingford, UK, p. 1–13.CrossRefGoogle Scholar
- Hodgson, J., 1990. Grazing management: Science in practise. Longman Scientific & Technical, UK.Google Scholar
- Hoffman, R., J. Wilson, D. Kronfeld, W. Cooper, L. Lawrence, D. Sklan and P. Harris, 2001. Hydrolyzable carbohydrates in pasture, hay, and horse feeds: Direct assay and seasonal variation. J. Anim. Sci. 79, 500–506.PubMedGoogle Scholar
- Hopkins, A., 2000. Herbage production. In: Hopkins, A. (ed.) Grass — its production and utilization. 3rd ed. Blackwell Science Ltd, Oxford, UK, p. 90–110.Google Scholar
- Housley, T.L. and J.J. Volenec, 1988. Fructan content and synthesis in leaf tissues of festuca-arundinacea. Plant Physiol. 86, 1247–1251.PubMedCrossRefGoogle Scholar
- Huhtanen, P., J. Nousiainen and M. Rinne, 2005. Prediction of silage composition and organic matter digestibility from herbage composition and pepsin-cellulase solubility. Agric. Food Sci. Finland 14, 154–165.CrossRefGoogle Scholar
- Huhtanen, P., J. Nousiainen and M. Rinne, 2006. Recent developments in forage evaluation with special reference to practical applications. Agric. Food Sci. Finland 15, 293–323.CrossRefGoogle Scholar
- Hume, D.E., 1991. Effect of cutting on production and tillering in prairie grass (bromus-willdenowii kunth) compared with 2 ryegrass (lolium) species. 1. Vegetative plants. Ann. Botany 67, 533–541.Google Scholar
- Jarvis S.C., 1993. Nitrogen cycling and losses from dairy farms. Soil Use and Management 9, 99–105.CrossRefGoogle Scholar
- Jones, M.B. and A. Lazenby (eds.), 1988. The grass crop: the physiological basis of production. Chapman and Hall Ltd, London, UK.Google Scholar
- Jung, H-J., D. Samac and G Sarath, 2012. Modifying crops to increase cell wall digestibility. Plant Sci. 185–186, 65–77.PubMedCrossRefGoogle Scholar
- Kagan, I.A., B.H. Kirch and J.R. Strickland, 2011. A chromatographic survey of methods for extracting long-chain grass fructans. Grass Forage Sci. 66, 434–448.CrossRefGoogle Scholar
- Kephart, K.D., D.R. Buxton and S.E. Taylor, 1992. Growth of C3 and C4 perennial grasses under reduced irradiance. Crop Sci. 32, 1033–1038.CrossRefGoogle Scholar
- Kuoppala, K., M. Rinne, J. Nousiainen and P. Huhtanen, 2008. The effect of cutting time of grass silage in primary growth and regrowth and the interactions between silage quality and concentrate level on milk production of dairy cows. Livestock Sci. 116, 171–182.CrossRefGoogle Scholar
- Lattanzi, F.A., 2010. C3/C4 grassland and climate change. In: H. Schnyder (editor) Grassland in a changing world: Proceedings of the 23th general meeting of the European Grassland Federation, Kiel, Germany. August 29th — September 2nd 2010. Grassland Science in Europe 15:3–13.Google Scholar
- Leafe, E.L., 1988. Introduction — the history of improved grasslands. In: Jones, M.B. and A. Lazenby (eds.) The grass crop. 1st ed. Chapman and Hall Ltd, London, UK, p. 1–23.CrossRefGoogle Scholar
- Lemaire, G. and D. Chapman, 1996. Tissue flows in grazed plant communities. In: Hodgson, J. and A.W. Illius (eds.) The ecology and management of grazing systems. CAB International, Wallingford, UK, p. 3–36.Google Scholar
- Lemaire, G., 1988. Sward dynamics under different management programmes. p. 7–12. In: Proceedings of the 12th general meeting of the European Grassland Federation, July 1988.Google Scholar
- Li, X., J. Weng and C. Chapple, 2008. Improvement of biomass through lignin modification. Plant J. 54, 569–581.PubMedCrossRefGoogle Scholar
- Longland, A. and B. Byrd, 2006. Pasture nonstructural carbohydrates and equine laminitis. J. Nutr. 136,2099S-2102S.PubMedGoogle Scholar
- Lybbert, T., P. Gibbs, N. Cohen, B. Scott and D. Sigler, 2007. Feeding alfalfa hay to exercising horses reduces the severity of gastric squamous mucosal ulceration. Proc. Amer. Assoc. Equine Practnr 53, 525–526.Google Scholar
- Martin-Rosset, W. and J.P. Dulphy, 1987. Digestibility interactions between forages and concentrates in horses: influence of feeding level - comparison with sheep. Livest. Prod. 17, 263–276.CrossRefGoogle Scholar
- Mayland, H.F., P.R. Cheeke, W. Majak and J.P. Goff, 2007. Forage-induced animal disorders. In: Barnes, R.F., C.J. Nelson, K.J. Moore and M. Collins (eds.) Forages: the science of grassland agriculture. 6th ed. Blackwell Publishing, Oxford, UK, p. 687–707.Google Scholar
- McDonald, P., A.R. Henderson and S.J.E. Heron, 1991. The biochemistry of silage 2nd ed. Chalcombe Publications, Bucks, UK. 340 p.Google Scholar
- Moorby, J., R. Evans, N. Scollan, J. Macraet and M. Theodorou, 2006. Increased concentration of water-soluble carbohydrate in perennial ryegrass (Lolium perenne L.). evaluation in dairy cows in early lactation. Grass Forage Sci. 61, 52–59.CrossRefGoogle Scholar
- Morin, C., G.F. Tremblay, G. Belanger, A. Bertrand, Y. Castonguay, R. Drapeau, R. Michaud, R. Berthiaume and G. Allard, 2012. Nonstructural carbohydrate concentration during field wilting of PM- and AM-cut alfalfa. Agron J. 104, 649–660.CrossRefGoogle Scholar
- Morrison, I., 1980. Changes in the lignin and hemicellulose concentrations of 10 varieties of temperate grasses with increasing maturity. Grass Forage Sci. 35, 287–293.CrossRefGoogle Scholar
- Morvan-Bertrand, A., J. Boucaud, J. Le Saos and M. Prud’homme, 2001. Roles of the fructans from leaf sheaths and from the elongating leaf bases in the regrowth following defoliation of Lolium perenne L. Planta 213, 109–120.PubMedCrossRefGoogle Scholar
- Moser, L.E. and J.A. Jennings, 2007. Grass and legume structure and morphology. In: Barnes, R.F., C.J. Nelson, K.J. Moore and M. Collins (eds.), Forages. The science of grassland agriculture. 6th ed., vol II. Blackwell publishing, Ames, Iowa, USA, p. 15–35.Google Scholar
- Müller, C., 2007. Wrapped forages for horses. Doctoral Thesis. Faculty of veterinary medicine and animal science, Swedish University of Agricultural Sciences, 57 p.Google Scholar
- Müller, C.E., D. Von Rosen and P. Uden, 2008. Effect of forage conservation method on microbial flora and fermentation pattern in forage and in equine colon and faeces. Livestock Sci. 119, 116–128.CrossRefGoogle Scholar
- Nadeau, J., F. Andrews, A. Mathew, R. Argenzio, J. Blackford, M. Sohtell and A. Saxton, 2000. Evaluation of diet as a cause of gastric ulcers in horses. Am. J. Vet. Res. 61, 784–790.PubMedCrossRefGoogle Scholar
- Nelson, C.J., 2000. Shoot morphological plasticity of grasses: Leaf growth vs. tillering. In: Lemaire, G., J. Hodgson, A. de Moraes, C. Nabinger and P.C. de F. Carvalho (eds.) Grassland ecophysiology and grazing ecology. CAB International, Wallingford, Oxon, UK, p. 101–126.CrossRefGoogle Scholar
- Ourry, A., J. Macduff and H. Ougham, 1996. The relationship between mobilization of N reserves and changes in translatable messages following defoliation in Lolium temulentum L and Lolium perenne L. J. Exp. Bot. 47, 739–747.CrossRefGoogle Scholar
- Pagan, J.D., 2009. Forages: The foundation for equine gastrointestinal health. Adv. Equine Nutr. 4, 17–24.Google Scholar
- Parsons, A.J., 1988. The effects of season and management on the growth of grass swards. In: Jones, M.B. and A. Lazenby (eds.), The grass crop. 1st ed. Chapman and Hall Ltd, London, UK, p. 129–177.CrossRefGoogle Scholar
- Pelletier, S., G.F. Tremblay, A. Bertrand, G. Belanger, Y. Castonguay and R. Michaud, 2010b. Drying procedures affect non-structural carbohydrates and other nutritive value attributes in forage samples. Anim. Feed Sci. Technol. 157, 139–150.CrossRefGoogle Scholar
- Pelletier, S., G.F. Tremblay, G. Belanger, A. Bertrand, Y. Castonguay, D. Pageau and R. Drapeau, 2010a. Forage nonstructural carbohydrates and nutritive value as affected by time of cutting and species. Agron. J. 102, 1388–1398.CrossRefGoogle Scholar
- Pitman, W.D., E.C. Holt, B.E. Conrad and E.C. Bashaw, 1983. Histological differences in moisture-stressed and nonstressed kleingrass forage. Crop Sci. 23, 793–795.CrossRefGoogle Scholar
- Pollock, C.J. and T. Jones, 1979. Seasonal patterns of fructan metabolism in forage grasses. New Phytol. 83, 9–15.CrossRefGoogle Scholar
- Ragnarsson, S. and A. Jansson, 2010. Comparison of grass haylage digestibility and metabolic plasma profile in Icelandic and Standardbred horses. J. Anim. Physiol. Nutr. 95, 273–279.CrossRefGoogle Scholar
- Ragnarsson, S. and J.E. Lindberg, 2008. Nutritional value of timothy haylage in Icelandic horses. Livest Sci. 113, 202–208.CrossRefGoogle Scholar
- Ragnarsson, S. and J.E. Lindberg, 2010. Nutritional value of mixed grass haylage in Icelandic horses. Livest Sci. 131, 83–87CrossRefGoogle Scholar
- Ralston, S.L., 2010. Nutrition: Horses In: Kahn, C.M. and S. Line (eds.) Merck Veterinary Manual. Merck & Co., Inc., Whitehouse Station, New Jersey, USA. Available at: http://www.merckvetmanual.com/mvm/index.jsp?cfile=htm/ bc/182602.htm.Google Scholar
- Reddy, M.S.S., F. Chen, G.L. Shadle, L. Jackson, H. Aljoe and R.A. Dixon, 2005. Targetted down-regulation of cytochrome P450 enzymes for forage quality improvement in alfalfa (Medicago sativa L.). PNAS 102, 16573–16578.PubMedCrossRefGoogle Scholar
- Reid, D. and N.H. Strachan, 1974. The effects of a wide range of nitrogen rates on some chemical constituents of herbage from perennial ryegrass swards with and without white clover. J. Agr. Sci., Cambridge 83, 393–401.Google Scholar
- Richards, J.H. and M.M. Caldwell, 1985. Soluble carbohydrates, concurrent photosynthesis and efficiency in regrowth following defoliation, a field-study with agropyron species. J. Appl. Ecol. 22, 907–920.CrossRefGoogle Scholar
- Richards, J.H., 1993. Physiology of plants recovering from defoliation. p. 85–94. In: Physiology of plants recovering from defoliation. Proceedings of the XVII International Grassland Congress, Palmerston North, New Zealand. 8–21 Feb. 1993.Google Scholar
- Rinne, M. and A. Nykanen, 2000. Timing of primary growth harvest affects the yield and nutritive value of timothy-red clover mixtures. Agric. Food Sci. Finland 9, 121–134.Google Scholar
- Robson, M.J., G.J.A. Ryle and J. Woledge, 1988. The grass plant - its for and function. In: Jones, M.B. and A. Lazenby (eds.) The grass crop. The physiological basis of production. Chapman & Hall, London, UK, p. 25–84.Google Scholar
- Roche, J.R., P. Dillon, S. Crosse and M. Rath, 1996. The effect of closing date of pasture in autumn and turnout date in spring on sward characteristics, dry matter yield and milk production of spring-calving dairy cows. Ir. J. Agric. and Food Res. 35, 127–140.Google Scholar
- Romera, A.J., D.G. McCall, J.M. Lee and M.G. Agnusdei, 2009. Improving the McCall herbage growth model. N. Z. J. Agric. Res. 52, 477–494.CrossRefGoogle Scholar
- Saastamoinen, M., 1994. Winter feeding of horses with preserved feeds and some by-products. a review. Livest. Prod. Sci. 40, p. 80.Google Scholar
- Saastamoinen, M.T. and M. Hellämäki, 2012. Forage analyses as a base of feeding of horses. In: Saastamoinen, M.T., M.J. Fradinho, N. Miraglia, and A.S. Santos (eds.) Forages and grazing in horse nutrition. Wageningen Academic Publishers, Wageningen, the Netherlands, p. 305–314.CrossRefGoogle Scholar
- Saikkonen, K., M. Helander and S.H. Faeth, 2004. Fungal endophytes: Hitch-hikers of the green world. In: Gillings, M. and A. Holmes (eds.) Plant microbiology. Garland Science/BIOS Scientific publishers, Oxford, UK, p. 77–95.Google Scholar
- Sanderson, M.A. and W.F. Wedin, 1989. Phenological stage and herbage quality relationships in temperature grasses and legumes. Agron. J. 81, 864–869.CrossRefGoogle Scholar
- Sanderson, M.A., D.W. Stair and M.A. Hussey, 1997. Physiological and morphological responses of perennial forages to stress. In: Sparks, D.L. (ed.) Advances in agronomy. 59th ed. Academic press, San Diego, California, USA.Google Scholar
- Särkijärvi, S., O. Niemeläinen, R. Sormunen-Cristian, and M. Saastamoinen, 2010. Suitability of grass species on equine pasture: Water soluble carbohydrates and grass preferences by horses. Grassland Science in Europe 15, 1000–1002.Google Scholar
- Särkijärvi, S., R. Sormunen-Cristian, T. Heikkilä, M. Rinne and M. Saastamoinen, 2012. Effect of grass species and cutting time on in vivo digestibility of silage by horses and sheep. Livestock Sci. 144, 230–239.CrossRefGoogle Scholar
- Sayers, H. and C. Mayne, 2001. Effect of early turnout to grass in spring on dairy cow performance. Grass Forage Sci. 56, 259–267.CrossRefGoogle Scholar
- Schnyder, H., R. Schäufele, R. de Visser and C.J. Nelson, 2000. An integrated view of C and N uses in leaf growth zones of defoliated grasses. In: Lemaire, G., J. Hodgson, A. de Moraes, P.C. de F. Carvalho and C. Nabinger (eds.) Grassland ecophysiology and grazing ecology. CABI Publishing, Wallingford, UK, p. 41–60.CrossRefGoogle Scholar
- Schwarz, A.G. and J.T. Reaney, 1989. Perennating structures and freezing tolerance of northern and southern populations of C4 grasses. Bot. Gaz. 150, 239–246.CrossRefGoogle Scholar
- Seppänen, M.M., K. Pakarinen, V. Jokela, J.R. Andersen, A. Fiil, A. Santanen and P. Virkajärvi, 2010. Vernalization response of Phleum pratense and its relationships to stem lignification and floral transition. Ann. Botany 106, 697–707.CrossRefGoogle Scholar
- Sheaffer, C.C., P.R. Peterson, M.H. Hall and J.B. Stordahl, 1992. Drought effects on yield and quality of perennial grasses in the north central united-states. J. Prod. Agric. 5, 556–561.Google Scholar
- Shewmaker, G.E., H.F. Mayland, C.A. Roberts, P. A. Harrison, N.J. Chatterton and D.A. Sleper, 2006. Daily carbohydrate accumulation in eight tall fescue cultivars. Grass Forage Sci. 61, 413–421.CrossRefGoogle Scholar
- Skinner, R.H. and K.J. Moore, 2007. Growth and development of forage plants. In: Barnes, R.F., C.J. Nelson, K.J. Moore and M. Collins (eds.) Forages: the science of grassland agriculture. 6th ed. Blackwell Publishing, Ames, Iowa, USA, p. 53–66.Google Scholar
- Smit, H.J., S. Tamminga and A. Elgersma, 2006. Dairy cattle grazing preference among six cultivars of perennial ryegrass. Agron. J. 98, 1213–1220.CrossRefGoogle Scholar
- Smith, D., 1967. Carbohydrates in grasses. II. Sugar and fructosan composition of the stem bases of bromegrass and timothy at several growth stages and in different plant parts at anthesis. Crop Sci. 7, 62–67.CrossRefGoogle Scholar
- Smith, D., 1972. Carbohydrate reserves of grasses. In: Youngner, V.B. and C.M. McKell (eds.) The biology and utilization of grasses. Academic Press, New York, NY, USA, p. 318–333.Google Scholar
- Sollenberger, L.E., R.A. Reis, L.G. Nussio, C.G. Chambliss and W.E. Kunkle, 2004. Conserved forage. In: Moser, L.E. (ed.) Warm-season (C4) grasses. Monograph 45. ASA-CSSA-SSSA, Madison, WI, USA, p. 355–387.Google Scholar
- Southwood, L.L., D.L. Evans, W.L. Bryden and R.J. Rose, 1993. Feeding practices in thoroughbred and standardbred racehorse stables. Aust. Vet. J. 70, 184–185.PubMedCrossRefGoogle Scholar
- Spollen, W.G. and C.J. Nelson, 1994. Response of fructan to water-deficit in growing leaves of tall fescue. Plant Physiol. 106, 329–336.PubMedGoogle Scholar
- Staniar, W.B., J.R. Bussard,, N.M. Repard, M.H. Hall and A.O. Burk, 2010. Voluntary intake and digestibility of teff gay fed to horses. J. Anim. Sci. 88, 3296–3303.PubMedCrossRefGoogle Scholar
- Suzuki, M., 1989. Fructans in forage grasses with varying degrees of coldhardiness. J. Plant Physiol. 134, 224–231.CrossRefGoogle Scholar
- Taiz, L. and E. Zeiger (eds.), 2010. Plant physiology. 5th ed. Sinauer Associates Inc., Sunderland, MA, USA.Google Scholar
- Taweel, H., B. Tas, H. Smit, A. Elgersma, J. Dijkstra and S. Tamminga, 2006. Grazing behaviour, intake, rumen function and milk production of dairy cows offered Lolium perenne containing different levels of water-soluble carbohydrates RID G-7680-2011. Livestock Sci. 102, 33–41.CrossRefGoogle Scholar
- Terry, R.A. and J.M.A. Tilley, 1964. The digestibility of the leaves and stems of perennial ryegrass, cocksfoot, timothy, tall fescue, lucerne and sainfoin, as measured by an in vitro procedure. Grass Forage Sci. 19, 363–372.CrossRefGoogle Scholar
- Thorsteinsson, B., P. Harrison and N. Chatterton, 2002. Fructan and total carbohydrate accumulation in leaves of two cultivars of timothy (Phleum pratense vega and climax) as affected by temperature. J. Plant Physiol. 159, 999–1003.CrossRefGoogle Scholar
- Thorvaldsson, G. and B. Fagerberg, 1988. Effects of weather on nutritional-value and phenological development of timothy. Swed. J. Agric. Res. 18, 51–59.Google Scholar
- Tomomatsu, H., 1994. Health-effects of oligosaccharides. Food Technol. 48, 61–65.Google Scholar
- Tremblay, G.F., G. Belanger and R. Drapeau, 2005. Nitrogen fertilizer application and developmental stage affect silage quality of timothy (Phleum pratense L.). Grass Forage Sci., 337–355.Google Scholar
- Uotila, R., T. Thuneberg and M. Saastamoinen, 2012. The usage of forage analyses in optimizing horse nutrition in Finland. In: Saastamoinen, M.T., M.J. Fradinho, N. Miraglia, and A.S. Santos (eds.) Forages and grazing in horse nutrition. Wageningen Academic Publishers, Wageningen, the Netherlands, p. 331–334.CrossRefGoogle Scholar
- Välisalo, M., S. Jaakkola, E. Saarisalo, T. Heikkilä and M. Seppänen, 2010. The sugar content of grasses in relation to horse feeding. Proceedings of Maataloustieteen Päivät 2010. www.smts.fi (In Finnish).
- Van Soest, P.J., 1994. Nutritional ecology of the ruminant. 2nd edition. Comstock Publishing Associates, Cornell University Press, Ithaca and London. 476 p.Google Scholar
- Virkajärvi, P. and K. Jarvenranta, 2001. Leaf dynamics of timothy and meadow fescue under Nordic conditions. Grass Forage Sci. 56, 294–304.CrossRefGoogle Scholar
- Virkajärvi, P., A. Sairanen, J. Nousiainen and H. Khalili, 2003. Sward and milk production response to early turnout of dairy cows to pasture in Finland. Agric. Food Sci. Finland 12, 21–34.Google Scholar
- Virkajärvi, P., K. Pakarinen, M. Hyrkäs, M. Seppanen and G. Belanger, 2012a. Tiller characteristics of timothy and tall fescue in relation to herbage mass accumulation. Crop Sci. 52, 1–11.CrossRefGoogle Scholar
- Virkajärvi, P., M. Hyrkäs, K. Pakarinen and R. Suomela, 2012b. Importance of senescence and dead material on nutritive value of grass silage. Proceedings of the XVI International Silage Conference, Hameenlinna, Finland. (in Press).Google Scholar
- Volaire, F. and F. Lelievre, 1997. Production, persistence, and water-soluble carbohydrate accumulation in 21 contrasting populations of Dactylis glomerata L. subjected to severe drought in the south of france. Aust. J. Agric. Res. 48, 933–944.CrossRefGoogle Scholar
- Volaire, F. and J. Gandoin, 1996. The effect of age of the sward on the relationship between water-soluble carbohydrate accumulation and drought survival in two contrasted populations of cocksfoot (Dactylis glomerata L). Grass Forage Sci. 51, 190–198.CrossRefGoogle Scholar
- Volenec, J., A. Ourry and B. Joern, 1996. A role for nitrogen reserves in forage regrowth and stress tolerance. Physiol. Plantarum 97, 185–193.CrossRefGoogle Scholar
- Volenec, J.J., 1986. Nonstructural carbohydrates in stem base components of tall fescue during regrowth. Crop Sci. 26, 122–127.CrossRefGoogle Scholar
- Volenec, J.J., and C.J. Nelson, 2007. Physiology of forage plants. In: Barnes, R.F., C.J. Nelson, K.J. Moore and M. Collins (eds.), Forages: The science of grassland agriculture. 6th ed. Blackwell Publishing, Ames, Iowa, USA, p. 37–52.Google Scholar
- Watts, K., 2005. A review of unlikely sources of excess carbohydrate in equine diets. J. Equine Vet. Sci. 25, 338–344.CrossRefGoogle Scholar
- Watts, K.A., 2009. Carbohydrates in forage: What is a safe grass? Adv. Equine Nutr. IV.Google Scholar
- Whitehead, D.C., 1995. Grassland nitrogen. CAB International, Wallingford, UK.Google Scholar
- Wilkins, P. and M. Humphreys, 2003. Progress in breeding perennial forage grasses for temperate agriculture. J. Agric. Sci. 140, 129–150.CrossRefGoogle Scholar
- Wilson, J.R. and P.M. Kennedy, 1996. Plant and animal constraints to voluntary feed intake associated with fibre characteristics and particle breakdown and passage in ruminants. Aust. J. Agric. Res. 47, 199–225.CrossRefGoogle Scholar
- Wilson, J.R., B. Deinum and F.M. Engels, 1991. Temperature effects on anatomy and digestibility of leaf and stem of tropical and temperate forage species. Neth. J. Agric. Sci. 39, 31–48.Google Scholar
- Woodward, S., 1998. Quantifying different causes of leaf and tiller death in grazed perennial ryegrass swards. N. Z. J. Agric. Res. 41, 149–159.CrossRefGoogle Scholar
- Wu, Z., 2004. Quantification of photoperiodic effects on growth of Phleum pratense. Doctor Scientiarum These 2006. 6. Agricultural University of Norway, Ås. Norway.Google Scholar
- Zhao, D., C.T. MacKown, P.J. Starks and B.K. Kindiger, 2008. Interspecies variation of forage nutritive value and nonstructural carbohydrates in perennial cool-season grasses. Agron. J. 100, 837–844.CrossRefGoogle Scholar