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Dynamics of forage ingestion, oral processing and digesta outflow from the rumen: a development in a mechanistic model of a grazing ruminant, MINDY

Published online by Cambridge University Press:  19 November 2018

P. Gregorini*
Affiliation:
Feed and Farm Systems Group, DairyNZ Ltd, Private Bag 3221, Hamilton 3240, New Zealand
F. D. Provenza
Affiliation:
Department of Wildland Resources, Utah State University, Logan 84322-5230, USA
J. J. Villalba
Affiliation:
Department of Wildland Resources, Utah State University, Logan 84322-5230, USA
P. C. Beukes
Affiliation:
Feed and Farm Systems Group, DairyNZ Ltd, Private Bag 3221, Hamilton 3240, New Zealand
M. J. Forbes
Affiliation:
Institute of Integrative and Comparative Biology, Faculty of Biological Sciences, 13 University of Leeds, LS2 9JT, UK
*
Author for correspondence: P. Gregorini, present address: Faculty of Agricultural and Life Sciences Lincoln University Lincoln 7647, Christchurch, New Zealand. E-mail: pablo.gregorini@lincoln.ac.nz

Abstract

Detailed representation of ingesta inflow to and digesta outflow from the rumen is critical for improving the modelling of rumen function and herbage intake of grazing ruminants. The objective of the current work was to extend a mechanistic model of a grazing ruminant, MINDY, to simulate the dynamic links between ingestive and digestive processes as affected by forage and sward features (e.g. sward structure, herbage chemical composition) as well as the internal state of the animal. The work integrates existing aspects of forage ingestion, oral physiology and rumen digestion that influence ingesta characteristics and digesta outflows from the rumen, respectively. The paper describes the structure and function of the new development, assessing the new model in terms of dynamic changes of oral processing of ingesta and rumen dilution rate under different grazing contexts. MINDY reproduces characteristics of ingesta inflow to and digesta outflow from the rumen of grazing ruminants, achieving temporal patterns of occurrence within and between meals, similar to those for grazing animals reported in the literature. The model realistically simulates changes in particle size distribution of the ingestive bolus, bolus weight and rumen dilution rate in response to contrasting grazing management regimes. The new concepts encoded in MINDY capture the underlying biological mechanisms that drive the dynamic link between ingestion and digestion patterns. This development advances in the understanding and modelling of grazing and digestive behaviour patterns of free-ranging ruminants.

Type
Modelling Animal Systems Research Paper
Copyright
Copyright © Cambridge University Press 2018 

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References

Acosta, A, Boudon, A and Peyraud, J-L (2007) Species and variety of herbage affects release of cell contents during ingestive mastication in dairy cows fed indoors. Animal Feed Science and Technology 132, 2848.Google Scholar
Argyle, JL and Baldwin, RL (1988) Modeling of rumen water kinetics and effects of rumen pH changes. Journal of Dairy Science 71, 11781188.Google Scholar
Bailey, CB and Balch, CC (1961) Saliva secretion and its relation to feeding in cattle. 2. The composition and rate of secretion of mixed saliva in the cow during rest. British Journal of Nutrition 15, 383402.Google Scholar
Bailey, AT, Erdman, R, Smith, L and Sharma, B (1990) Particle size reduction during initial mastication of forages by dairy cattle. Journal of Animal Science 68, 20842094.Google Scholar
Bailey, DW, Gross, JE, Laca, EA, Rittenhouse, LR, Coughenour, MB, Swift, DM and Sims, PL (1996) Mechanisms that result in large herbivore grazing distribution patterns. Journal of Range Management 49, 386400.Google Scholar
Balch, CC (1958) Observations on the act of eating in cattle. British Journal of Nutrition 12, 330345.Google Scholar
Balch, CC and Campling, RC (1962) Regulation of voluntary food intake in ruminants. Nutrition Abstracts and Reviews 32, 669686.Google Scholar
Baldwin, RL (1995) Modeling Ruminant Digestion and Metabolism. London, UK: Chapman and Hall.Google Scholar
Baldwin, RL, Thornley, JHM and Beever, DE (1987) Metabolism of the lactating cow: II. Digestive elements of a mechanistic model. Journal of Dairy Research 54, 107131.Google Scholar
Barboza, PS and Bowyer, RT (2000) Sexual segregation in dimorphic deer: a new gastrocentric hypothesis. Journal of Mammalogy 81, 473489.Google Scholar
Baumont, R, Cohen-Salmon, D, Prache, S and Sauvant, D (2004) A mechanistic model of intake and grazing behaviour in sheep integrating sward architecture and animal decisions. Animal Feed Science and Technology 112, 528.Google Scholar
Bergman, CM, Fryxell, JM, Gates, CC and Fortin, D (2001) Ungulate foraging strategies: energy maximizing or time minimizing? Journal of Animal Ecology 70, 289300.Google Scholar
Boudon, A, Acosta, A, Delagarde, R and Peyraud, JL (2006) Release of cell contents and comminution of particles of perennial ryegrass herbage during ingestion by dairy cows fed indoors or grazing. Grass and Forage Science 61, 205217.Google Scholar
Bruce, LA and Huber, TL (1973) Inhibitory effect of acid in the intestine on rumen motility in sheep. Journal of Animal Science 37, 164168.Google Scholar
Burns, JC and Sollenberger, LE (2002) Grazing behavior of ruminants and daily performance from warm-season grasses. Crop Science 42, 873881.Google Scholar
Carvalho, PCF, Bremm, B, Mezzalira, JC, Fonseca, L, da Trindade, JK, Bonnet, O, Tischler, M, Genro, TCM, Nabinger, C and Laca, EA (2015) Can animal performance be predicted from short-term grazing processes? Animal Production Science 55, 319327.Google Scholar
Chacon, E and Stobbs, TH (1976) Influence of progressive defoliation of a grass sward on the eating behavior of cattle. Australian Journal of Agricultural Research 27, 709727.Google Scholar
Chilibroste, P (1999) Grazing time, the missing link. PhD, Wageningen University and Research Centre.Google Scholar
Chilibroste, P, Soca, P, Mattiauda, DA, Bentancur, O and Robinson, PH (2007) Short term fasting as a tool to design effective grazing strategies for lactating dairy cattle: a review. Australian Journal of Experimental Agriculture 47, 10751084.Google Scholar
Chilibroste, P, Dijkstra, J, Robinson, PH and Tamminga, S (2008) A simulation model “CTR Dairy” to predict the supply of nutrients in dairy cows managed under discontinuous feeding patterns. Animal Feed Science and Technology 143, 148173.Google Scholar
Chilibroste, P, Gibb, MJ, Soca, P and Mattiauda, DA (2015) Behavioral adaptation of dairy cows to changes in feeding management: do they follow a predictable pattern? Animal Production Science 55, 328338.Google Scholar
Clauss, M and Lechner-Doll, M (2001) Differences in selective reticulo-ruminal particle retention as a key factor in ruminant diversification. Oecologia 129, 321327.Google Scholar
Clauss, M, Hummel, J and Streich, WJ (2006) The dissociation of the fluid and particle phase in the forestomach as a physiological characteristic of large grazing ruminants: an evaluation of available, comparable ruminant passage data. European Journal of Wildlife Research 52, 8898.Google Scholar
Clauss, M, Hume, ID and Hummel, J (2010) Evolutionary adaptations of ruminants and their potential relevance for modern production systems. Animal: An International Journal of Animal Bioscience 4, 979992.Google Scholar
Clauss, M, Lechner, I, Barboza, P, Collins, W, Tervoort, TA, Südekum, K-H, Codron, D and Hummel, J (2011) The effect of size and density on the mean retention time of particles in the reticulorumen of cattle (Bos primigenius f. Taurus), muskoxen (Ovibos moschatus) and moose (Alces alces). British Journal of Nutrition 105, 634644.Google Scholar
Coffey, KP, Paterson, JA, Saul, CS, Coffey, LS, Turner, KE and Bowman, JG (1989) The influence of pregnancy and source of supplemental protein on intake, digestive kinetics and amino acid absorption by ewes. Journal of Animal Science 67, 18051814.Google Scholar
Demment, MW and Greenwood, GB (1988) Forage ingestion: effects of sward characteristics and body size. Journal of Animal Science 66, 23802392.Google Scholar
Demment, MW and Laca, EA (1994) Reductionism and synthesis in the grazing sciences: models and experiments. Proceedings of the Australian Soiecty of Animal Production 20, 616.Google Scholar
Deswysen, AG, Ellis, WC and Pond, KR (1987) Interrelationships among voluntary intake, eating and ruminating behavior and ruminal motility of heifers fed corn silage. Journal of Animal Science 64, 835841.Google Scholar
Dewhurst, RJ, Davies, DR and Merry, RJ (2000) Microbial protein supply from the rumen. Animal Feed Science and Technology 85, 121.Google Scholar
Dijkstra, J, Oenema, O and Bannink, A (2011) Dietary strategies to reducing N excretion from cattle: implications for methane emissions. Current Opinion in Environmental Sustainability 3, 414422.Google Scholar
Dijkstra, J, Oenema, O, Van Groenigen, JW, Spek, JW, Van Vuuren, AM and Bannink, A (2013) Diet effects on urine composition of cattle and N2O emissions. Animal: An International Journal of Animal Bioscience 7, 292302.Google Scholar
Dove, H and Milne, JA (1994) Digesta flow and rumen microbial protein production in ewes grazing perennial ryegrass. Australian Journal of Agricultural Research 45, 12291245.Google Scholar
Dove, H, Milne, JA, Sibbald, AM, Lamb, CS and McCormack, HA (1988) Circadian variation in abomasal digesta flow in grazing ewes during lactation. British Journal of Nutrition 60, 653668.Google Scholar
Drescher, MF (2003) Grasping Complex Matter: Large Herbivore Foraging in Patches of Heterogeneous Resources. Wageningen, the Netherlands: Wageningen Universiteit.Google Scholar
Faichney, G (2005) Digesta flow. In Forbes, JM and France, J (eds), Quantitative Aspects of Ruminant Digestion and Metabolism. Wallingford, UK: CAB International, pp. 5385.Google Scholar
Forbes, JM and Gregorini, P (2015) The catastrophe of meal eating. Animal Production Science 55, 350359.Google Scholar
Fortin, D, Fryxell, JM and Pilote, R (2002) The temporal scale of foraging decisions in bison. Ecology 83, 970982.Google Scholar
Freer, M and Campling, RC (1965) Factors affecting the voluntary intake of food by cows. 7. The behaviour and reticular motility of cows given diets of hay, dried grass, concentrates and ground, pelleted hay. British Journal of Nutrition 19, 195207.Google Scholar
Gill, J, Campling, RC and Westgarth, DR (1966) A study of chewing during eating in the cow. British Journal of Nutrition 20, 1323.Google Scholar
Gill, M, Robinson, PH and Kennelly, JJ (1999) Diurnal patterns in rumen volume and composition of digesta flowing into the duodenum. Animal Science 69, 237249.Google Scholar
Greenwood, GB and Demment, MW (1988) The effect of fasting on short-term cattle grazing behaviour. Grass and Forage Science 43, 377386.Google Scholar
Gregorini, P (2011) Estado interno. Estímulos que motivan el consumo y ciertas conductas ingestivas de rumiantes en pastoreo. In Cangiano, CA and Brizuela, MA (eds), Produccion Animal en Pastoreo. Buenos Aires, Argentina: INTA, pp. 291320.Google Scholar
Gregorini, P (2012) Diurnal grazing pattern: its physiological basis and strategic management. Animal Production Science 52, 416430.Google Scholar
Gregorini, P, Gunter, SA, Masino, CA and Beck, PA (2007) Effects of ruminal fill on short-term herbage intake rate and grazing dynamics of beef heifers. Grass and Forage Science 62, 346354.Google Scholar
Gregorini, P, Gunter, SA and Beck, PA (2008) Matching plant and animal processes to alter nutrient supply in strip-grazed cattle: timing of herbage and fasting allocation. Journal of Animal Science 86, 10061020.Google Scholar
Gregorini, P, Gunter, SA, Beck, PA, Caldwell, J, Bowman, MT and Coblentz, WK (2009 a) Short-term foraging dynamics of cattle grazing swards with different canopy structures. Journal of Animal Science 87, 38173824.Google Scholar
Gregorini, P, Soder, KJ and Kensinger, RS (2009 b) Effects of rumen fill on short-term ingestive behavior and circulating concentrations of ghrelin, insulin, and glucose of dairy cows foraging vegetative micro-swards. Journal of Dairy Science 92, 20952105.Google Scholar
Gregorini, P, Gunter, SA, Bowman, MT, Caldwell, JD, Masino, CA, Coblentz, WK and Beck, PA (2011) Effect of herbage depletion on short-term foraging dynamics and diet quality of steers grazing wheat pastures. Journal of Animal Science 89, 38243830.Google Scholar
Gregorini, P, Beukes, PC, Romera, AJ, Levy, G and Hanigan, MD (2013) A model of diurnal grazing patterns and herbage intake of a dairy cow, MINDY: model description. Ecological Modelling 270, 1129.Google Scholar
Gregorini, P, Beukes, P, Waghorn, G, Pacheco, D and Hanigan, M (2015 a) Development of an improved representation of rumen digesta outflow in a mechanistic and dynamic model of a dairy cow, Molly. Ecological Modelling 313, 293306.Google Scholar
Gregorini, P, Villalba, JJ, Provenza, FD, Beukes, PC and Forbes, JM (2015 b) Modelling preference and diet selection patterns by grazing ruminants: a development in a mechanistic model of a grazing dairy cow, MINDY. Animal Production Science 55, 360375.Google Scholar
Gregorini, P, Beukes, PC, Dalley, D and Romera, AJ (2016) Screening for diets that reduce urinary nitrogen excretion and methane emissions while maintaining or increasing production by dairy cows. Science of the Total Environment 551–552, 3241.Google Scholar
Gregorini, P, Villalba, JJ, Chilibroste, P and Provenza, FD (2017) Grazing management: setting the table, designing the menu and influencing the diner. Animal Production Science 57, 12481268.Google Scholar
Gregorini, P, Provenza, FD, Villalba, JJ, Beukes, PC and Forbes, MJ (2018) Diurnal patterns of urination and drinking by grazing ruminants: a development in a mechanistic model of a grazing ruminant, MINDY. Journal of Agricultural Science, Cambridge 156, 7181.Google Scholar
Gunter, SA, Judkins, MB, Krysl, LJ, Broesder, JT, Barton, RK, Rueda, BR, Hallford, DM and Holcombe, DW (1990) Digesta kinetics, ruminal fermentation characteristics and serum metabolites of pregnant and lactating ewes fed chopped alfalfa hay. Journal of Animal Science 68, 38213831.Google Scholar
Hanigan, MD, Bateman, HG, Fadel, JG, McNamara, JP and Smith, NE (2006) An ingredient-based input scheme for Molly. In Kebreab, E, Dijkstra, J, Bannink, A, Gerrits, WJJ and France, J (eds), Nutrient Digestion and Utilization in Farm Animals: Modelling Approaches. Walingford, UK: CAB International, pp. 328348.Google Scholar
Hanigan, MD, Appuhamy, JADRN and Gregorini, P (2013) Revised digestive parameter estimates for the Molly cow model. Journal of Dairy Science 96, 38673885.Google Scholar
Hanks, DR, Judkins, MB, McCracken, BA, Holcombe, DW, Krysl, LJ and Park, KK (1993) Effects of pregnancy on digesta kinetics and ruminal fermentation in beef cows. Journal of Animal Science 71, 28092814.Google Scholar
Hayes, MR, Moore, RL, Shah, SM and Covasa, M (2004) 5-HT3 receptors participate in CCK-induced suppression of food intake by delaying gastric emptying. American Journal of Physiology: Regulatory, Integrative and Comparative Physiology 287, R817R823.Google Scholar
Hogan, JP, Keeney, PA and Weston, RH (1985) Factors affecting the intake of feed by grazing animals. In Wheeler, JL, Pearson, CJ and Robards, GE (eds), Temperate Pastures: Their Production, Use and Management. Canberra, Australia: CSIRO, pp. 317327.Google Scholar
Hutchings, JB and Lillford, P (1988) The perception of food texture: the philosophy of the breakdown path. Journal of Texture Studies 19, 103115.Google Scholar
Illius, AW and Gordon, IJ (1987) The allometry of food intake in grazing ruminants. Journal of Animal Ecology 56, 989999.Google Scholar
Illius, AW and Gordon, IJ (1992) Modelling the nutritional ecology of ungulate herbivores: evolution of body size and competitive interactions. Oecologia 89, 428434.Google Scholar
Jung, HG and Allen, MS (1995) Characteristics of plant cell walls affecting intake and digestibility of forages by ruminants. Journal of Animal Science 73, 27742790.Google Scholar
Kennedy, PM (2005) Particle dynamics. In Dijkman, J, Forbes, JM and France, J (eds), Quantitative Aspects of Ruminant Digestion and Metabolism. Wallingford, UK: CAB International, pp. 123156.Google Scholar
Kennedy, PM and Murphy, MR (1988) The nutritional implications of differential passage of particles through the ruminant alimentary tract. Nutrition Research Reviews 1, 189208.Google Scholar
Laca, EA and Demment, MW (1991) Herbivory: the dilemma of foraging in a spatially heterogeneous food environment. In Palo, RT and Robbins, CT (eds), Plant Defenses Against Mammalian Herbivory. Boca Raton, FL, USA: CRC Press, pp. 3044.Google Scholar
Laca, EA, Ungar, ED, Seligman, N and Demment, MW (1992) Effects of sward height and bulk density on bite dimensions of cattle grazing homogeneous swards. Grass and Forage Science 47, 91102.Google Scholar
Laca, EA, Ungar, ED and Demment, MW (1994) Mechanisms of handling time and intake rate of a large mammalian grazer. Applied Animal Behavior Science 39, 319.Google Scholar
Lechner, I, Barboza, P, Collins, W, Fritz, J, Günther, D, Hattendorf, B, Hummel, J, Südekum, K-H and Clauss, M (2010) Differential passage of fluids and different-sized particles in fistulated oxen (Bos primigenius f. Taurus), muskoxen (Ovibos moschatus), reindeer (Rangifer tarandus) and moose (Alces alces): rumen particle size discrimination is independent from contents stratification. Comparative Biochemistry and Physiology, Part A: Molecular and Integrative Physiology 155, 211222.Google Scholar
Li, Y, Wu, XY and Owyang, C (2004) Serotonin and cholecystokinin synergistically stimulate rat vagal primary afferent neurons. Journal of Physiology 559, 651662.Google Scholar
López, S, Hovell, FDD and MacLeod, NA (1994) Osmotic pressure, water kinetics and volatile fatty acid absorption in the rumen of sheep sustained by intragastric infusions. British Journal of Nutrition 71, 153168.Google Scholar
Lucas, PW, Prinz, JF, Agrawal, KR and Bruce, IC (2002) Food physics and oral physiology. Food Quality and Preference 13, 203213.Google Scholar
Luginbuhl, J-M, Pond, KR, Burns, JC and Russ, JC (1989) Effects of ingestive mastication on particle dimensions and weight distribution of coastal bermudagrass hay fed to steers at four levels. Journal of Animal Science 67, 538546.Google Scholar
McLeod, MN and Minson, DJ (1988) Large particle breakdown by cattle eating ryegrass and alfalfa. Journal of Animal Science 66, 992999.Google Scholar
Meng, Q, Kerley, MS, Ludden, PA and Belyea, RL (1999) Fermentation substrate and dilution rate interact to affect microbial growth and efficiency. Journal of Animal Science 77, 206214.Google Scholar
Moseley, G and Jones, JR (1984) The physical digestion of perennial ryegrass (Lolium perenne) and white clover (Trifolium repens) in the foregut of sheep. British Journal of Nutrition 52, 381390.Google Scholar
Okine, EK, Mathison, GW and Hardin, RT (1989) Effects of changes in frequency of reticular contractions on fluid and particulate passage rates in cattle. Journal of Animal Science 67, 33883396.Google Scholar
Owens, FN, Secrist, DS, Hill, WJ and Gill, DR (1998) Acidosis in cattle: a review. Journal of Animal Science 76, 275286.Google Scholar
Pérez-Barbería, FJ and Gordon, IJ (1998) Factors affecting food comminution during chewing in ruminants: a review. Biological Journal of the Linnean Society 63, 233256.Google Scholar
Pérez-Prieto, LA and Delagarde, R (2012) Meta-analysis of the effect of pregrazing pasture mass on pasture intake, milk production, and grazing behavior of dairy cows strip-grazing temperate grasslands. Journal of Dairy Science 95, 53175330.Google Scholar
Pérez-Prieto, LA and Delagarde, R (2013) Meta-analysis of the effect of pasture allowance on pasture intake, milk production, and grazing behavior of dairy cows grazing temperate grasslands. Journal of Dairy Science 96, 66716689.Google Scholar
Pittroff, W and Soca, P (2006) Physiology and models of feeding behaviour and intake regulation in ruminants. In Bels, V (ed.), Feeding in Domestic Vertebrates: from Structure to Behaviour. Wallingford, UK: CAB International, pp. 278301.Google Scholar
Pond, KR, Tolley, EA, Ellis, WC and Matis, JH (1984) A method for describing the weight distribution of particles from sieved forage. In Kennedy, PM (ed.), Techniques in Particle Size Analysis of Feed and Digesta in Ruminants. Edmonton, AB, Canada: Canadian Society of Animal Science, pp. 123133.Google Scholar
Pond, KR, Ellis, WC, Lascano, CE and Akin, DE (1987) Fragmentation and flow of grazed coastal bermudagrass through the digestive tract of cattle. Journal of Animal Science 65, 609618.Google Scholar
Poppi, DP, Minson, DJ and Ternouth, JH (1981) Studies of cattle and sheep eating leaf and stem fractions of grasses. 2. Factors controlling the retention of feed in the reticulo-rumen. Australian Journal of Agricultural Research 32, 109121.Google Scholar
Poppi, DP, France, J and McLennan, SR (2000) Intake, passage and digestibility. In Theodorou, MK and France, J (eds), Feeding Systems and Feed Evaluation Models. Wallingford: CAB International, pp. 3553.Google Scholar
Prinz, JF and Lucas, PW (1997) An optimization model for mastication and swallowing in mammals. Proceedings of the Royal Society of London. Series B: Biological Sciences 264, 17151721.Google Scholar
Rykiel, EJ (1996) Testing ecological models: the meaning of validation. Ecological Modelling 90, 229244.Google Scholar
Saunders, PT (1980) An Introduction to Catastrophe Theory. Cambridge, UK: Cambridge University Press.Google Scholar
Sauvant, D, Baumont, R and Faverdin, P (1996) Development of a mechanistic model of intake and chewing activities of sheep. Journal of Animal Science 74, 27852802.Google Scholar
Schettini, MA, Prigge, EC and Nestor, EL (1999) Influence of mass and volume of ruminal contents on voluntary intake and digesta passage of a forage diet in steers. Journal of Animal Science 77, 18961904.Google Scholar
Seo, S, Lanzas, C, Tedeschi, LO and Fox, DG (2007) Development of a mechanistic model to represent the dynamics of liquid flow out of the rumen and to predict the rate of passage of liquid in dairy cattle. Journal of Dairy Science 90, 840855.Google Scholar
Seo, S, Lanzas, C, Tedeschi, LO, Pell, AN and Fox, DG (2009) Development of a mechanistic model to represent the dynamics of particle flow out of the rumen and to predict rate of passage of forage particles in dairy cattle. Journal of Dairy Science 92, 39814000.Google Scholar
Shipley, LA, Gross, JE, Spalinger, DE, Hobbs, NT and Wunder, BA (1994) The scaling of intake rate in mammalian herbivores. American Naturalist 143, 10551082.Google Scholar
Spalinger, DE, Robbins, CT and Hanley, TA (1986) The assessment of handling time in ruminants: the effect of plant chemical and physical structure on the rate of breakdown of plant particles in the rumen of mule deer and elk. Canadian Journal of Zoology 64, 312321.Google Scholar
Stuth, JW and Angell, RF (1982) Effect of seasonal herbage allowance on bolus weights of cattle. Journal of Range Management 35, 163165.Google Scholar
Taweel, HZ, Tas, BM, Dijkstra, J and Tamminga, S (2004) Intake regulation and grazing behavior of dairy cows under continuous stocking. Journal of Dairy Science 87, 34173427.Google Scholar
Thiago, LR, Gill, M and Sissons, JW (1992) Studies of method of conserving grass herbage and frequency of feeding in cattle. 2. Eating behavior, rumen motility and rumen fill. British Journal of Nutrition 67, 319336.Google Scholar
Ulyatt, MJ (1983) Plant fibre and regulation of digestion in the ruminant. In Wallace, G and Bell, L (eds), Fibre in Human and Animal Nutrition. Wellington, New Zealand: Royal Society of New Zealand, pp. 103107.Google Scholar
Ulyatt, MJ, Waghorn, GC, John, A, Reid, CSW and Monro, J (1984) Effect of intake and feeding frequency on feeding behavior and quantitative aspects of digestion in sheep fed chaffed lucerne hay. Journal of Agricultural Science, Cambridge 102, 645657.Google Scholar
Vanzant, ES, Cochran, RC and Johnson, DE (1991) Pregnancy and lactation in beef heifers grazing tallgrass prairie in the winter: influence on intake, forage utilization, and grazing behavior. Journal of Animal Science 69, 30273038.Google Scholar
Wade, MH and Carvalho, PCF (2000) Defoliation patterns and herbage intake on pastures. In Lemaire, G, Hodgson, J, Moraes, A, Nabinger, C and Carvalho, PCF (eds), Grassland Ecophysiology and Grazing Ecology. Wallingsford, UK: CAB International, pp. 233248.Google Scholar
Waite, R (1963) Grazing behaviour. In Worden, AN, Sellers, KC and Tribe, DE (eds), Animal Health, Production and Pasture. London, UK: Longman, pp. 286309.Google Scholar
Waldo, DR, Smith, LW and Cox, EL (1972) Model of cellulose disappearance from the rumen. Journal of Dairy Science 55, 125129.Google Scholar
Wilson, JR and Kennedy, PM (1996) Plant and animal constraints to voluntary feed intake associated with fibre characteristics and particle breakdown and passage in ruminants. Australian Journal of Agricultural Research 47, 199225.Google Scholar
Wilson, JR and Mertens, DR (1995) Cell wall accessibility and cell structure limitations to microbial digestion of forage. Crop Science 35, 251259.Google Scholar