Register      Login
Animal Production Science Animal Production Science Society
Food, fibre and pharmaceuticals from animals
RESEARCH ARTICLE

Diurnal grazing pattern: its physiological basis and strategic management

P. Gregorini
+ Author Affiliations
- Author Affiliations

DairyNZ, Private Bag 3221, Hamilton, New Zealand. Email: pablo.gregorini@dairynz.co.nz

Animal Production Science 52(7) 416-430 https://doi.org/10.1071/AN11250
Submitted: 21 October 2011  Accepted: 2 February 2012   Published: 3 April 2012

Abstract

Within a day, grazing decisions such as ‘when’ to begin, ‘which’ frequency and ‘how’ to distribute grazing events determine ruminants’ diurnal grazing pattern. Ruminants can have between three and five daily grazing events. The major grazing events occur in the early morning and late afternoon/early evening; the later grazing event is the longest and most significant in terms of herbage intake. This review first attempts to answer ‘why does this happen?’ and then to examine evidence for managing this pattern to improve animal production. Due to photosynthesis and transpiration during the day, herbage accumulates DM, sugars and essential fatty acids, which dilute fibre and protein contents and facilitate herbage particle breakdown during ingestion. Diurnal fluctuations in light intensity stimulate circadian release of neuropetides and hormones, providing the cue to start grazing and modulating ingestive-digestive behaviours that interact with the diurnal fluctuation in herbage feeding value. Grazing decisions depend on grazing environments, the current state of the animal, and on past and anticipated states of the animal. The dusk grazing event seems to be an adaptative feeding strategy to maximise daily energy acquisition, providing a steady release of nutrients over night. Hunger deceives ruminants and makes them graze at dawn, when herbage presents the lowest feeding value. Hunger, however, can be used to concentrate and intensify grazing events. Strategic management of these interactions emerges as the tool to alter the frequency, intensity and temporal distribution of diurnal grazing events, and thereby to increase and modulate nutrient supply to and productivity of grazing ruminants.

Additional keywords: behaviour, herbage intake, nutrient supply, ruminants.


References

Abrahamse PA, Tamminga S, Dijkstra J (2009) Effect of daily movement of dairy cattle to fresh grass in morning or afternoon on intake, grazing behaviour, rumen fermentation and milk production. The Journal of Agricultural Science 147, 721–730.
Effect of daily movement of dairy cattle to fresh grass in morning or afternoon on intake, grazing behaviour, rumen fermentation and milk production.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BD1MXhtlant7fF&md5=134bcad3438c3432a4b07b3c008ce50aCAS |

Aikman PC, Mills JAN, Reynolds CK, Crompton LA (2011) Effect of feeding frequency and protein supplementation on methane production by Holstein cows. Journal of Dairy Science 94, E-Suppl. 1, 607

Akabayashi A, Levin N, Paez X, Alexander JT, Leibowitz SF (1994) Hypothalamic neuropeptide Y and its gene expression: relation to light dark cycle and circulating corticosterone. Molecular and Cellular Neurosciences 5, 210–218.
Hypothalamic neuropeptide Y and its gene expression: relation to light dark cycle and circulating corticosterone.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DyaK2cXkt1Ohtbk%3D&md5=93422ef9d3dd0abdcf1777c62aeede91CAS |

Allen MS, Bradford BJ, Harvatine KJ (2005) The cow as a model to study food intake regulation. Annual Review of Nutrition 25, 523–547.
The cow as a model to study food intake regulation.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BD2MXpvF2mt7s%3D&md5=d6289d84025e0fc8d2d036f8907ac432CAS |

Andueza D Delgado I Munoz F 2011 Variation of digestibility and intake by sheep of lucerne (Medicago saviva L.) hay cuts at sunrise or sunset. Journal of Agricultural Science Cambridge 10.1071/S00218596111000542

Bailey D, Gross W, Laca EA, Rittenhouse LR, Coughenour MB, Swift DM, Sims PL (1996) Mechanisms that result in large herbivore grazing distribution patterns. Journal of Range Management 49, 386–400.

Berthelot XM, Lautentie JP, Revault J, Ferney J, Toutain PL (1990) Circadian profile and production rate of melatonin in the cow. Domestic Animal Endocrinology 7, 315–322.
Circadian profile and production rate of melatonin in the cow.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DyaK3cXlsVGksr0%3D&md5=31075c8f439671ca0be4eeed27106693CAS |

Borghese A, Barile VL, Tergaro GM, Pilla AM, Parmeggani A (1994) Melatonin trend during seasons in heifers and buffalo cows. In ‘Proceedings of fourth buffalo congress. Vol. 3, Sao Paulo, Brazil, 27–30 June’. pp. 528–530.

Brainard GC, Hanifin JP, Greeson JM, Byrne B, Glickman G, Gerner E, Rollag M (2001) Action spectrum for melatonin regulation in humans: evidence for a novel circadian photoreceptor. The Journal of Neuroscience 21, 6405–6414.

Brito AF, Tremblay GF, Lapierre H, Bertrand A, Castonquay Y, Belanger G, Benchaar C, Oullet DR, Berthiaime R (2008) Alfalfa cut at sundown and harvested as baleage improves milk yield of late-lactation dairy cows. Journal of Dairy Science 91, 3968–3982.
Alfalfa cut at sundown and harvested as baleage improves milk yield of late-lactation dairy cows.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BD1cXht1SrurvE&md5=ffdb91d78f5a75fdec2502e4958dbce6CAS |

Brito AF, Tremblay GF, Lapierre H, Bertrand A, Castonquay Y, Belanger G, Benchaar C, Oullet DR, Berthiaime R (2009) Alfalfa cut at sundown and harvested as baleage increased bacterial protein synthesis in late-lactation dairy cows. Journal of Dairy Science 92, 1092–1107.
Alfalfa cut at sundown and harvested as baleage increased bacterial protein synthesis in late-lactation dairy cows.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BD1MXivVaktLc%3D&md5=ea6f797d15887d7b336913c16973184bCAS |

Browse JP, Roughan G, Slack CR (1981) Light control of fatty acid synthesis and diurnal fluctuation of fatty acid composition in leaves. Biochemistry Journal 196, 347–354.

Bruce LA, Huber TL (1973) Inhibitory of acid in the intestine on rumen motility in sheep. Journal of Animal Science 37, 164–168.

Burritt EA, Mayland HF, Provenza FD, Miller RL, Burns JC (2005) Effect of added sugar on preference and intake by sheep of hay cut in the morning versus the afternoon. Applied Animal Behaviour Science 94, 245–254.
Effect of added sugar on preference and intake by sheep of hay cut in the morning versus the afternoon.Crossref | GoogleScholarGoogle Scholar |

Casler MD, Scheider DK, Combs DK (1996) Development and application of a selection criterion for particle size breakdown of smooth bromegrass leaves. Animal Feed Science and Technology 61, 57–71.
Development and application of a selection criterion for particle size breakdown of smooth bromegrass leaves.Crossref | GoogleScholarGoogle Scholar |

Chapman DF, Parsons AJ, Cosgrove GP, Baker DJ, Marotti DM, Venning KJ, Rutter SM, Hill J, Thomson AN (2007) Impact of spatial patterns in pasture on animal grazing behavior, intake and performance. Crop Science 47, 399–415.
Impact of spatial patterns in pasture on animal grazing behavior, intake and performance.Crossref | GoogleScholarGoogle Scholar |

Charnov EL, Gordon HO, Hyatt K (1976) Ecological implications of resource depression. American Naturalist 110, 247–259.
Ecological implications of resource depression.Crossref | GoogleScholarGoogle Scholar |

Chilibroste P, Tamminga S, Boer H, Gibb MJ, den Dikken G (2000) Duration of regrowth of ryegrass (Lolium perenne). Effects on grazing behavior, intake, rumen fill, and fermentation of lactating dairy cows. Journal of Dairy Science 83, 984–995.
Duration of regrowth of ryegrass (Lolium perenne). Effects on grazing behavior, intake, rumen fill, and fermentation of lactating dairy cows.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BD3cXjsVWksro%3D&md5=a7052886ca81fcfdb45a148ff5308bd9CAS |

Chilibroste P, Mattiauda DA, Elizondo F, Coster A (2004) Herbage allowance and grazing session allocation in dairy cows. Effect on milk production and composition. In ‘Proceeding of the 2nd International Symposium of Grassland Ecophysiology and Grazing Ecology, Curitiba, Brazil’. (Eds A Moraes, E Pizarro, PC Carvalho, S Carneiro da Silva, JR Dittrich), p. 25.

Chilibroste P, Gibb MJ, Tamminga S (2005) Pasture characteristics and animal performance. In ‘Quantitative aspects of ruminant digestion and metabolism’. (Eds J France, M Forbes, J Dijkstra) pp. 681–706. (CAB International: Wallingford)

Chilibroste P, Soca P, Mattiauda DA, Bentancur O, 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, 1075–1084.
Short-term fasting as a tool to design effective grazing strategies for lactating dairy cattle: a review.Crossref | GoogleScholarGoogle Scholar |

Ciavarella TA, Simpson RJ, Dove H, Leury BJ, Sims IM (2000) Diurnal changes in the concentration of water-soluble carbohydrates in Phalaris aquatica L. pasture in spring, and the effect of short-term shading. Australian Journal of Agricultural Research 51, 749–756.
Diurnal changes in the concentration of water-soluble carbohydrates in Phalaris aquatica L. pasture in spring, and the effect of short-term shading.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BD3cXmtFKjtrY%3D&md5=0ca090bab838c56b98db372854979f19CAS |

Clark CEF, McLeod KLM, Glassey CB, Gregorini P, Betteridge K, Jago JG (2010) Capturing urine while maintaining pasture intake, milk production, and animal welfare of dairy cows in early and late lactation. Journal of Dairy Science 93, 2280–2286.
Capturing urine while maintaining pasture intake, milk production, and animal welfare of dairy cows in early and late lactation.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BC3cXnvVansLs%3D&md5=083f7908f123f4227971c29af0fca6d6CAS |

Clissold FJ (2008) The biomechanics of chewing and plant fracture: mechanism and implications. Advances in Insect Physiology 34, 317–372.
The biomechanics of chewing and plant fracture: mechanism and implications.Crossref | GoogleScholarGoogle Scholar |

Conrad HR, Baile CA, Mayer J (1977) Changing meal patterns and suppression of feed intake with increasing amounts of dietary non protein nitrogen in ruminants. Journal of Dairy Science 60, 1725–1733.
Changing meal patterns and suppression of feed intake with increasing amounts of dietary non protein nitrogen in ruminants.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DyaE1cXlvFWrtA%3D%3D&md5=4f2a79b15d7dea66017ec801d4462a3cCAS |

Cosgrove GP, Edwards GR (2007) Control of grazing intake. In ‘Pasture and supplements for grazing animals’. (Eds PV Rattray, IM Brooks, AM Nicol) pp. 61–80. (New Zealand Society of Animal Production: Hamilton)

Cosgrove GP, Waghorn GC, Parsons AJ (1999) Exploring the nutritional basis of preference and diet selection by sheep. Proceedings of the New Zealand Grassland Association 61, 175–180.

Crompton LA, Mills JAN, Reynolds CK France J (2009) Fluctuation in methane emission in response to feeding pattern to dairy cows. In ‘Proceedings of the 7th International Workshop in Modelling Nutrient Digestion and Utilization in Farm Animals, Paris, France 10–12 Sept. 2009’. (Eds DE Dalley, JR Roche, PJ Moate) p. 28.

Cruickshank GJ (1986) Nutritional constraints to lamb growth at pasture. PhD Thesis, Lincoln University, New Zealand.

Dalley DE, Roche JR, Moate PJ, Grainger C (2001) More frequent allocation of herbage does not improve the milk production of dairy cows in early lactation. Australian Journal of Experimental Agriculture 41, 593–599.
More frequent allocation of herbage does not improve the milk production of dairy cows in early lactation.Crossref | GoogleScholarGoogle Scholar |

Delagarde R, Peyraud JL, Delaby L, Faverdin P (2000) Vertical distribution of biomass, chemical composition, and pepsin-cellulase digestibility in a perennial ryegrass sward: interaction with month and year, re-growth age and time of day. Animal Feed Science and Technology 84, 49–68.
Vertical distribution of biomass, chemical composition, and pepsin-cellulase digestibility in a perennial ryegrass sward: interaction with month and year, re-growth age and time of day.Crossref | GoogleScholarGoogle Scholar |

Deswysen A, Ehrlein HJ (1979) Radiography of intake and (pseudo) rumination behaviour. Annals of Veterinary Research 10, 208–210.

Di Marco OE, Aello MS, Mendez DG (1996) Energy expenditure of cattle grazing of pasture of low and high availability. Animal Science (Penicuik, Scotland) 63, 45–50.
Energy expenditure of cattle grazing of pasture of low and high availability.Crossref | GoogleScholarGoogle Scholar |

Dougherty CT, Cornelius PL, Bradley NW, Lauriault LM (1989) Ingestive behaviour of beef heifers within grazing session. Applied Animal Behaviour Science 23, 341–351.
Ingestive behaviour of beef heifers within grazing session.Crossref | GoogleScholarGoogle Scholar |

Dove H, Milne JA, Sibbald AM, Lamb CS, McCormack HA (1988) Circadian variation in abomasal digesta flow in grazing ewes during lactation. The British Journal of Nutrition 60, 653–658.
Circadian variation in abomasal digesta flow in grazing ewes during lactation.Crossref | GoogleScholarGoogle Scholar | 1:STN:280:DyaL1M7hvVWquw%3D%3D&md5=5b264d1535f90acda86ce56d3bdfc6d4CAS |

Fernstrom JD (1988) Tryptophan, serotonin and carbohydrate appetite: will the real carbohydrate craver stand up! The Journal of Nutrition 118, 1417–1419.

Fisher DS, Mayland HF, Burns JC (1999) Variation in ruminants’ preference for tall fescue hays cut either at sundown or at sunup. Journal of Animal Science 77, 762–768.

Fisher DS, Mayland HF, Burns JC (2002) Variation in ruminant preference for alfalfa hays cut at sunup and sundown. Crop Science 42, 231–237.
Variation in ruminant preference for alfalfa hays cut at sunup and sundown.Crossref | GoogleScholarGoogle Scholar |

Fisher DS, Burns JC, Mayland HF (2005) Ruminant selection among switchgrass hays cut at either sundown or sunup. Crop Science 45, 1394–1402.
Ruminant selection among switchgrass hays cut at either sundown or sunup.Crossref | GoogleScholarGoogle Scholar |

Freer M, 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. The British Journal of Nutrition 19, 195–207.
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.Crossref | GoogleScholarGoogle Scholar | 1:STN:280:DyaF2M%2FpsFCntg%3D%3D&md5=a7056eb718e908cb364dd7401a7489ceCAS |

Geary N (2004) Endocrine control of eating. CCK, leptin and ghrelin. Physiology & Behavior 81, 719–733.
Endocrine control of eating. CCK, leptin and ghrelin.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BD2cXlsVWjtr8%3D&md5=1bb4fe5e969b9c9f77736a75287c6c52CAS |

Gebbie FE, Forsyth IA, Arendt J (1999) Effects of maintaining solstice light and temperature on reproductive activity, coat growth, plasma prolactin and melatonin in goats. Journal of Reproduction and Fertility 116, 25–33.
Effects of maintaining solstice light and temperature on reproductive activity, coat growth, plasma prolactin and melatonin in goats.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DyaK1MXjvVemsLs%3D&md5=74c1e94ef4f595682994eab343bc42f9CAS |

Gekara OJ, Prigge EC, Bryan WB, Nestor EL, Seidel G (2005) Influence of sward height, daily timing of concentrate supplementation, and restricted time for grazing on forage utilization by lactating beef cows. Journal of Animal Science 83, 1435–1444.

Gibb MJ (1998) Animal grazing/intake terminology and definitions. In ‘Pasture ecology and animal intake. Proceedings of a workshop held in Dublin 1996. Concerned Action. AIR3–CT93–0947’. Occasional Publication 3. pp. 21–37.

Gibb MJ (2006) Grassland management with emphasis on grazing behaviour. In ‘Fresh herbage for dairy cattle’. (Eds A Elgersma, J Dijkstra, S Tamminga) pp. 141–157. (Springer: The Netherlands)

Gibb MJ, Huckle CA, Nuthall R (1998) Effect of time of day on grazing behaviour by lactating dairy cows. Grass and Forage Science 53, 41–46.
Effect of time of day on grazing behaviour by lactating dairy cows.Crossref | GoogleScholarGoogle Scholar |

Givens DI, Cottrill BR, Davies M, Lee PA, Mandsbridge RJ, Moss AR (2001) Sources of n-3 polyunsaturated fatty acids additional to fish oil for livestock diets: a review. Nutrition Abstracts and Review Series B 70, 3

Greenwood GB, Demment MW (1988) The effect of fasting on short-term cattle grazing behavior. Grass and Forage Science 43, 377–386.
The effect of fasting on short-term cattle grazing behavior.Crossref | GoogleScholarGoogle Scholar |

Gregorini P (2011) Internal state. Stimuli that motivates particular ingestive mechanisms and herbage intake in grazing ruminants. In ‘Producción animal en pastoreo’. (Eds C Cangiano, M Brizuela) pp. 514–565.

Gregorini P, Tamminga S, Gunter SA (2006a) Review: behavior and daily grazing patterns of cattle. The Professional Animal Scientist 22, 201–209.

Gregorini P, Eirin M, Refi R, Ursino M, Ansin O, Gunter SA (2006b) Timing of herbage allocation. Effect on beef heifers daily grazing pattern and performance. Journal of Animal Science 84, 1943–1950.
Timing of herbage allocation. Effect on beef heifers daily grazing pattern and performance.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BD28Xmt1aqtLk%3D&md5=e537b437e49e1d06864539d7183ce9d9CAS |

Gregorini P, Gunter SA, Masino CA, Beck PA (2007a) Effects of ruminal fill on intake rate and grazing dynamics of beef heifers. Grass and Forage Science 62, 346–354.
Effects of ruminal fill on intake rate and grazing dynamics of beef heifers.Crossref | GoogleScholarGoogle Scholar |

Gregorini P, Eirin M, Wade MH, Refi R, Ursino M, Ansin O, Gunter SA (2007b) The effects of a morning fasting on the evening grazing behavior and performance of strip-grazed beef heifers. The Professional Animal Scientist 23, 642–648.

Gregorini P, Eirin M, Agnelli L, Refi R, Ansin O, Masino CA, Wade MH, Soder K, Gunter SA (2007c) Diurnal eating pattern and performance of cattle strip grazed with afternoon herbage allocation or continuously stocked. In ‘Proceedings of the American Forage and Grassland Congress Annual Meeting, State College, PA’. p. 23.

Gregorini P, Gunter SA, Beck PA, Soder KJ, Tamminga S (2008a) REVIEW: the interaction of diurnal grazing pattern, ruminal metabolism, nutrient supply and management in cattle. The Professional Animal Scientist 24, 308–318.

Gregorini P, Gunter SA, Beck PA (2008b) Matching plant and animal processes to alter nutrient supply in strip grazed cattle: timing of herbage and fasting allocation. Journal of Animal Science 86, 1006–1020.
Matching plant and animal processes to alter nutrient supply in strip grazed cattle: timing of herbage and fasting allocation.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BD1cXjvFeru7k%3D&md5=d67b4e78bd3ce9ced23aa411cef871bfCAS |

Gregorini P, Soder KJ, Sanderson MA (2008c) CASE STUDY: a snapshot in time of fatty acids composition of grass herbage as affected by time of day. The Professional Animal Scientist 24, 675–680.

Gregorini P, Soder KJ, Sanderson MA, Ziegler G (2009a) Toughness, particle size and chemical composition of meadow fescue (Festuca pratensis Hud.) herbage as affected by time of day. Animal Feed Science and Technology 151, 330–336.
Toughness, particle size and chemical composition of meadow fescue (Festuca pratensis Hud.) herbage as affected by time of day.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BD1MXmtVGqs7o%3D&md5=fe0d65527fe45f79388f833dda298b54CAS |

Gregorini P, Soder KJ, Kensinger RS (2009b) Effect of rumen fill on foraging behavior, intake rate, and plasma ghrelin, serum insulin and glucose levels of cattle grazing a vegetative micro-sward. Journal of Dairy Science 92, 2095–2105.
Effect of rumen fill on foraging behavior, intake rate, and plasma ghrelin, serum insulin and glucose levels of cattle grazing a vegetative micro-sward.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BD1MXlsFymsrY%3D&md5=a4860f5e4cb56a62ca0292da68bf0e15CAS |

Gregorini P, Clark CEF, Jago JG, Glassey CB, McLeod K, Romera AJ (2009c) Restricting time at pasture: effects on dairy cow herbage intake, foraging behavior, hunger-related hormones and metabolite concentration during the first grazing session. Journal of Dairy Science 92, 4572–4580.
Restricting time at pasture: effects on dairy cow herbage intake, foraging behavior, hunger-related hormones and metabolite concentration during the first grazing session.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BD1MXhtVKqtrzF&md5=ae96e4141de683b670c120a6207ec57aCAS |

Gregorini P, Gunter SA, Beck PA, Caldwell J, Bowman MT, Coblentz WK (2009d) Short-term foraging dynamics of cattle grazing swards with different canopy structure. Journal of Animal Science 87, 3817–3824.
Short-term foraging dynamics of cattle grazing swards with different canopy structure.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BD1MXhtlentLbP&md5=b1f90db48b2315484abb0a97a4b0be6eCAS |

Gregorini P, Beukes P, Bryant R, Romera AJ (2010a) Invited review. Simulating the effect of feeding strategies on nitrogen excretions and methane emission from grazing dairy cows. In ‘Proceedings of the Australasian Dairy Symposium’. (Eds G Edwards, R Bryant) pp. 29–43. (Lincoln University: Lincoln, New Zealand)

Gregorini P, Soder KJ, Waghorn G (2010b) Effects of timing of corn silage supplementation on digestion, fermentation pattern and nutrient flow during continuous culture fermentation of a short and intensive orchardgrass meal. Journal of Dairy Science 93, 3722–3729.
Effects of timing of corn silage supplementation on digestion, fermentation pattern and nutrient flow during continuous culture fermentation of a short and intensive orchardgrass meal.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BC3cXht1Ggur7J&md5=61a2019dd6d78890f143d59a4c65f5bcCAS |

Gregorini P, Gunter SA, Bowman MT, Caldwell J, Masino CA, Coblentz WK, Beck PA (2011) Effect of herbage depletion on short-term foraging dynamics and diet quality of steers grazing wheat pastures. Journal of Animal Science
Effect of herbage depletion on short-term foraging dynamics and diet quality of steers grazing wheat pastures.Crossref | GoogleScholarGoogle Scholar |

Griggs TC, MacAdam JW, Mayland HF, Burns JC (2005) Nonstructural carbohydrate and digestibility patterns in orchardgrass swards during daily defoliation sequences initiated in evening and morning. Crop Science 45, 1295–1304.
Nonstructural carbohydrate and digestibility patterns in orchardgrass swards during daily defoliation sequences initiated in evening and morning.Crossref | GoogleScholarGoogle Scholar |

Hayes MR, Moore RL, Shah SM, Covasa M (2004) 5–HT3 receptors participate in CCK-induced suppression of food intake by delaying gastric empting. American Journal of Physiology Regulatory Integrative Physiology 287, 817–823.
5–HT3 receptors participate in CCK-induced suppression of food intake by delaying gastric empting.Crossref | GoogleScholarGoogle Scholar |

Herzog ED, Schwartz WJ (2002) A neural clockwork for encoding circadian time. Journal of Applied Physiology 92, 401–408.

Hoffmann K, Illenorva H, Vanacek J (1981) Effect of photoperiod of one minute light at night time in the pineal rhythm on N-acetyltrnaferasa activity in the D-jungarian hamster Phodppus singarus. Biology of Reproduction 24, 551–556.
Effect of photoperiod of one minute light at night time in the pineal rhythm on N-acetyltrnaferasa activity in the D-jungarian hamster Phodppus singarus.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DyaL3MXhvV2mur4%3D&md5=85e3e3b566105c3e5aad26db9b4a66fbCAS |

Hogan JP, Kenny PA, Weston RH (1987) Factor affecting the intake of feed by grazing animals. In ‘Temperate pastures: their production and management’. (Eds JL Wheeler, CJ Pearson, GE Robards) pp. 317–327. (Australian Wool Corporation/CSIRO: Melbourne)

Iason GR, Mantecon AR, Sim DA, Gonzalez J, Foreman E, Bermudez FF, Elston DA (1999) Can grazing sheep compensate for a daily foraging time constraints? Journal of Animal Ecology 68, 87–93.
Can grazing sheep compensate for a daily foraging time constraints?Crossref | GoogleScholarGoogle Scholar |

Janssen PH (2010) Influence of hydrogen on rumen methane formation and fermentation balances through microbial growth kinetics and fermentation thermodynamics. Animal Feed Science and Technology 160, 1–22.
Influence of hydrogen on rumen methane formation and fermentation balances through microbial growth kinetics and fermentation thermodynamics.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BC3cXhtV2itLvF&md5=b20b53db2aca14b3fd7d115c50c43895CAS |

Kennaway DJ, Frith RG, Phillipou G, Seamark RW (1983) Patterns of progesterone, melatonin and prolactin secretions in ewes maintaining in four different photoperiods. The Journal of Endocrinology 97, 229–242.
Patterns of progesterone, melatonin and prolactin secretions in ewes maintaining in four different photoperiods.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DyaL3sXhvFantLY%3D&md5=5840b11146c9263f2e74b3d0132d19f8CAS |

Kennedy PM (1985) Effect of rumination in reduction of particle size of digesta by cattle. Australian Journal of Agricultural Research 36, 819–828.
Effect of rumination in reduction of particle size of digesta by cattle.Crossref | GoogleScholarGoogle Scholar |

Kennedy E, O’Donovan M, McEvoy M, Murphy JP (2009) Effect of restricted access time to pasture on dairy cow milk production, grazing behavior and DM intake. Journal of Dairy Science 92, 168–176.
Effect of restricted access time to pasture on dairy cow milk production, grazing behavior and DM intake.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BD1MXlsVOqug%3D%3D&md5=c523f7e719c35632fd2560e6d944c922CAS |

Kollmann MT, Locher M, Hirche F, Eder K, Meyer HHD, Bruckmaier RM (2008) Effects of tryptophan supplementation on plasma tryptophan and related hormone levels in heifers and dairy cows. Domestic Animal Endocrinology 34, 14–24.
Effects of tryptophan supplementation on plasma tryptophan and related hormone levels in heifers and dairy cows.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BD2sXhtlWkt7jM&md5=7880241812deb43e1b3b9b21470eb785CAS |

Kolver ES, MacMillan KL (1993) Short term changes in selected metabolites in pasture fed dairy cows during peak lactation. Proceedings of the New Zealand Society of Animal Production 53, 77–81.

Kristensen T, Oudshoorn F, Munksgaard L, Soegaard K (2007) Effect of time at pasture combined with restricted indoor feeding on production and behaviour in dairy cows. Animal 1, 439–448.
Effect of time at pasture combined with restricted indoor feeding on production and behaviour in dairy cows.Crossref | GoogleScholarGoogle Scholar | 1:STN:280:DC%2BC38vptFKitA%3D%3D&md5=a869bf95bb21fe3180b51cc2d3a24cd4CAS |

Krysl LJ, Hess BW (1993) Influence of supplementation on grazing behaviour of cattle. Journal of Animal Science 71, 2546–2555.

Laca EA, Ungar ED, Demment MW (1994) Mechanisms of handling time and intake rate of a large mammalian grazer. Applied Animal Behaviour Science 39, 3–19.
Mechanisms of handling time and intake rate of a large mammalian grazer.Crossref | GoogleScholarGoogle Scholar |

Leibowitz SF (1992) Neurochemical-neuroendocrine systems in the brain controlling macronutrient intake and metabolism. Trends in Neurosciences 15, 491–497.
Neurochemical-neuroendocrine systems in the brain controlling macronutrient intake and metabolism.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DyaK3sXntlOnsQ%3D%3D&md5=b40ded8d410791820bc07cb3c5187e06CAS |

Li Y, Wu XY, Owyan C (2004) Serotonin and cholecystokinin synergistically stimulate rat vagal primary afferent neurons. The Journal of Physiology 559, 651–662.
Serotonin and cholecystokinin synergistically stimulate rat vagal primary afferent neurons.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BD2cXotVymu7g%3D&md5=b2f7d4bd2842870aa6a97f540970daaaCAS |

Linnane MI, Brereton AJ, Giller PS (2001) Seasonal changes in circadian grazing patterns of Kerry cows (Bos taurus) in semi-feral conditions in Killarney National Park, Co. Kerry, Ireland. Applied Animal Behaviour Science 71, 277–292.
Seasonal changes in circadian grazing patterns of Kerry cows (Bos taurus) in semi-feral conditions in Killarney National Park, Co. Kerry, Ireland.Crossref | GoogleScholarGoogle Scholar |

Loor JJ, Soriano FD, Lin X, Hirbein JH, Polan CE (2003) Grazing allowance after the morning or afternoon milking for lactating cows fed a total mixed ration (TMR) enhances trans11–18:1 and cis9,trans11–18:2 (rumenic acid) in milk fat to different extents. Animal Feed Science and Technology 109, 105–119.
Grazing allowance after the morning or afternoon milking for lactating cows fed a total mixed ration (TMR) enhances trans11–18:1 and cis9,trans11–18:2 (rumenic acid) in milk fat to different extents.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BD3sXntVGqt7o%3D&md5=87914f68fb2d79e82dc1fca07050c15cCAS |

Mattiauda DA, Tamminga S, Elizondo F, Gibb MJ, Chilibroste P (2004) Effect of allowance and timing of grazing session on dairy cows grazing permanent pastures. In ‘Proceeding of the 2nd International Symposium of Grassland Ecophysiology and Grazing Ecology, Curitiba, Brazil’. (Eds A Moraes, E Pizarro, PC Carvalho, S Carneiro da Silva, JR Dittrich).

Mayland HF, Shewmaker GE, Burns JC, Fisher D (1998) Morning and evening harvest effects on animal performance. In ‘Proceeding of the Alfalfa Symposium’. Reno, Nevada, USA.

McLeod MN, Minson DJ (1988) Large particle breakdown by cattle eating ryegrass and alfalfa. Journal of Animal Science 66, 992–999.

Meier S, Kolver EK, Verkerk GA, Roche JR (2010) Effect of divergent Holstein-Friesian strain and diet on diurnal patterns of plasma metabolites and hormones. The Journal of Dairy Research 77, 432–437.
Effect of divergent Holstein-Friesian strain and diet on diurnal patterns of plasma metabolites and hormones.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BC3cXht1aktL%2FE&md5=04a4c3d5d32cd152bea3e16dff3c6a63CAS |

Metz JHM (1975) Time patterns of feeding and rumination in domestic cattle. PhD Thesis, Wageningen Agricultural University, The Netherlands.

Miner JL (1992) Recent advances in central control of intake in ruminants. Journal of Animal Science 70, 1283–1289.

Newman JA, Parsons AJ, Thornley JHM, Penning PD, Krebs JR (1995) Optimal diet selection by a general herbivore. Functional Ecology 9, 255–268.
Optimal diet selection by a general herbivore.Crossref | GoogleScholarGoogle Scholar |

Nie Z, Ward G, Michael A (2001) Impact of pugging by dairy cows on pastures and indicators of pugging damage to pasture soil in south-western Victoria. Australian Journal of Agricultural Research 52, 37–44.
Impact of pugging by dairy cows on pastures and indicators of pugging damage to pasture soil in south-western Victoria.Crossref | GoogleScholarGoogle Scholar |

Nikkhah A (2011) Bioscience of ruminant intake evolution: feeding time models. Advances in Bioscience and Biotechnology 2, 271–274.
Bioscience of ruminant intake evolution: feeding time models.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BC38XhsVOjsbk%3D&md5=d2dd3bc9324d76921e9e927922a9bf1dCAS |

O’Brien JS, Benson AA (1964) Isolation and fatty acid composition of the plant sulfolipid and galactolipids. Journal of Lipid Research 5, 432–436.

Okine EK, Mathison GW, Hardin RT (1989) Effects of changes in frequency or reticular contractions on fluid and particle passage rates in cattle. Journal of Animal Science 67, 388–396.

Orr RJ, Penning PD, Harvey A, Champion RA (1997) Diurnal patterns of intake rate by sheep grazing monocultures of rye grass or white clover. Applied Animal Behaviour Science 52, 65–77.
Diurnal patterns of intake rate by sheep grazing monocultures of rye grass or white clover.Crossref | GoogleScholarGoogle Scholar |

Orr RJ, Rutter SM, Penning PD, Rook AJ (2001a) Matching grass supply to grazing patterns for dairy cows. Grass and Forage Science 56, 352–361.
Matching grass supply to grazing patterns for dairy cows.Crossref | GoogleScholarGoogle Scholar |

Orr RJ, Penning PD, Rutter SM, Champion RA, Harvey A, Rook AJ (2001b) Intake rate during meals and duration for sheep in different hunger states, grazing grass or white clover swards. Applied Animal Behaviour Science 75, 33–45.
Intake rate during meals and duration for sheep in different hunger states, grazing grass or white clover swards.Crossref | GoogleScholarGoogle Scholar |

Pande T, Valentine I, Betteridge K, Mackay A, Horne D (2000) Pasture damage and regrowth from cattle treading. In ‘Proceeding of the New Zealand Grassland Association’. pp. 155–160.

Penning PD, Parsons AJ, Orr RJ (1991) Patterns of ingestive behaviour of sheep continuously stocked on monocultures of ryegrass or white clover. Applied Animal Behaviour Science 31, 237–250.
Patterns of ingestive behaviour of sheep continuously stocked on monocultures of ryegrass or white clover.Crossref | GoogleScholarGoogle Scholar |

Perez-Barbería FJ, Gordon IJ (1998) Factors affecting food comminution during chewing in ruminants: a review. Biological Journal of the Linnean Society. Linnean Society of London 63, 233–256.

Pérez-Ramírez E, Delagarde R, Payreaud JL (2009) Restricting daily time at pasture at low and high herbage allowance: effects on herbage intake and behavioral adaptation of lactating dairy cows. Journal of Dairy Science 92, 3331–3340.
Restricting daily time at pasture at low and high herbage allowance: effects on herbage intake and behavioral adaptation of lactating dairy cows.Crossref | GoogleScholarGoogle Scholar |

Phillips CJC, Leaver JD (1986) Seasonal and diurnal variation of grazing behaviour of dairy cows. In ‘Proceedings of the British Grassland Society. Occasional Symposium No. 19’. pp. 98–104. (British Grassland Society: Hurley, UK)

Pittroff W, Soca P (2006) Physiology and models of feeding behavior and intake regulation in ruminants. In ‘Feeding in domestic vertebrates: from structure to behaviour’. pp. 278–301. (Ed. V Bels) (CAB International: Wallingford)

Roche JR, Sheahan AJ, Chagas LM, Boston RC (2008) Short communication: change in plasma ghrelin in dairy cows following an intravenous glucose challenge. Journal of Dairy Science 91, 1005–1010.
Short communication: change in plasma ghrelin in dairy cows following an intravenous glucose challenge.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BD1cXivFemt7s%3D&md5=296ef7b0af78cc99d552bf6e32835776CAS |

Roche JR, Blache D, Kay JK, Miller RD, Sheahan AJ, Miller W (2009) Neuroendocrine and physiological regulation of intake in domesticated ruminants: a review. Nutrition Research Reviews 21, 207–234.
Neuroendocrine and physiological regulation of intake in domesticated ruminants: a review.Crossref | GoogleScholarGoogle Scholar |

Rook AJ, Huckle CA (1997) Activity bout criteria for grazing dairy cows. Applied Animal Behaviour Science 54, 89–96.
Activity bout criteria for grazing dairy cows.Crossref | GoogleScholarGoogle Scholar |

Rook AJ, Huckle CA, Penning PD (1994) Effect of sward height and concentrate supplementation on the ingestive behaviour of spring calving dairy cows grazing grass clover swards. Applied Animal Behaviour Science 40, 101–112.
Effect of sward height and concentrate supplementation on the ingestive behaviour of spring calving dairy cows grazing grass clover swards.Crossref | GoogleScholarGoogle Scholar |

Rutter SM (2006) Diet preferences for grass and legumes in free-ranging domestic sheep and cattle: current theory and future application. Applied Animal Behaviour Science 97, 17–35.
Diet preferences for grass and legumes in free-ranging domestic sheep and cattle: current theory and future application.Crossref | GoogleScholarGoogle Scholar |

Rutter SM, Tainton V, Chamion RA, le Grice P (2002) The effect of a total solar eclipse on grazing behavior of dairy cattle. Applied Animal Behaviour Science 79, 273–283.
The effect of a total solar eclipse on grazing behavior of dairy cattle.Crossref | GoogleScholarGoogle Scholar |

Sarkar M, Prakash BS (2005) Circadian variation in plasma concentrations of melatonin and prolactin during breeding and non-breeding seasons in yak (Poephagus grunniens L.). Animal Reproduction Science 90, 149–162.
Circadian variation in plasma concentrations of melatonin and prolactin during breeding and non-breeding seasons in yak (Poephagus grunniens L.).Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BD2MXhtFKmt7rN&md5=96bd632fe5544d35108163e7f9f1fb55CAS |

Sauvé AK, Huntington GB, Whisnant CS, Burns JC (2010) Intake, digestibility, and nitrogen balance of steers fed gamagrass baleage top dressed at two rates of nitrogen and harvested at sunset and sunrise. Crop Science 50, 427–437.
Intake, digestibility, and nitrogen balance of steers fed gamagrass baleage top dressed at two rates of nitrogen and harvested at sunset and sunrise.Crossref | GoogleScholarGoogle Scholar |

Soca P (2000) Efecto del tiempo de pastoreo y nivel de suplementación sobre el consume, conducta y parámetros productivos d vacas lecheras MS Thesis, Universidad de Chile, Chile.

Soca P, Beretta V, Heinzen M, Bentancur O (2002) Effect of pasture height and control of grazing time on grazing behavior and defoliation dynamic of growing beef cattle. In ‘Symposium responding to the increasing global demand for animal products’. British Society Animal Science, 12–15 November 2002, Universidad Autónoma de Yucatán México.

Spalinger DE, Robbins CT, 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, 312–321.
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.Crossref | GoogleScholarGoogle Scholar |

Stobbs TH (1970) Automatic measurement of grazing time by dairy cows on tropical grass and legume pastures. Tropical Grasslands 4, 237–244.

Stobbs TH (1975) Factors limiting the nutritional value of grazed tropical pastures for beef and milk production. Tropical Grasslands 9, 141–150.

Stobbs TH (1977) Short-term effects of herbage allowance on milk production, milk composition and grazing time of cows grazing nitrogen-fertilized tropical grass pasture. Australian Journal of Experimental Agriculture and Animal Husbandry 17, 892–898.
Short-term effects of herbage allowance on milk production, milk composition and grazing time of cows grazing nitrogen-fertilized tropical grass pasture.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DyaE1cXnt1altw%3D%3D&md5=e0b67a2da0ec50894f3d2a1fcbd7241aCAS |

Strubbe JH, Woods CS (2004) The timing of meals. Psychological Review 111, 128–141.
The timing of meals.Crossref | GoogleScholarGoogle Scholar |

Sugino T, Hasegawa Y, Kurose Y, Kojima M, Kangawa K, Terashima Y (2004) Effects of ghrelin on food intake and neuroendocrine function in sheep. Animal Reproduction Science 82–83, 183–194.
Effects of ghrelin on food intake and neuroendocrine function in sheep.Crossref | GoogleScholarGoogle Scholar |

Swainson N, Martin C, Muetzel S, Pinares-Patiño C (2011) Hydrogen emissions from sheep: a spill-over for methanogenesis. In ‘Advances in Animal Biosciences. Proceedings of the 8th International Symposium on the Nutrition of Herbivores, Vol. 2, Issue 2’. p. 530.

Taweel HZ, Tas BM, Dijkstra J, Tamminga S (2004) Intake regulation and grazing behavior of dairy cows under continuous stocking. Journal of Dairy Science 87, 3417–3427.
Intake regulation and grazing behavior of dairy cows under continuous stocking.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BD2cXotFykurw%3D&md5=bcaa25524aafe9600f8381fda0312383CAS |

Thiago LRL (1988) Voluntary intake of forages by ruminants: factors related to eating behaviour and rumen fill. PhD Thesis, University of Reading.

Thomson BC, Cruickshank GJ, Poppi DP, Sykes AR (1985) Diurnal patterns of rumen fill in grazing sheep. Proceedings of the New Zealand Society of Animal Production 45, 117–112.

Toates F (2002) Physiology, motivation and the organization of behaviour. In ‘The ethology of domestic animals: an introductory text’. (Ed. P Jensen) pp. 31–50. (CAB International: Wallingford, UK)

Trevaskis LM, Fulkerson WJ, Gooden M (2001) Provision of certain carbohydrate-based supplements to pasture-fed sheep, as well as time of harvesting of the pasture, influences pH, ammonia concentration and microbial protein synthesis in the rumen. Australian Journal of Experimental Agriculture 41, 21–27.
Provision of certain carbohydrate-based supplements to pasture-fed sheep, as well as time of harvesting of the pasture, influences pH, ammonia concentration and microbial protein synthesis in the rumen.Crossref | GoogleScholarGoogle Scholar |

Trevaskis LM, Fulkerson WJ, Nandra KS (2004) Effect of time of feeding carbohydrate supplements and pasture on production of dairy cows. Livestock Production Science 85, 275–285.
Effect of time of feeding carbohydrate supplements and pasture on production of dairy cows.Crossref | GoogleScholarGoogle Scholar |

Vasta V, Pagano RI, Luciano G, Scerra M, Caparra P, Foti F, Cilione C, Biondi L, Priolo A, Avondo M (2012) Effect of morning vs. afternoon grazing on intramuscular fatty acid composition in lamb. Meat Science 90, 93–98.
Effect of morning vs. afternoon grazing on intramuscular fatty acid composition in lamb.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BC3MXht1ens7%2FL&md5=9fd50c95f6732dfcfdcb5c3ed16fdfeeCAS |

Vibart RE, Pacheco D, Lowe K, Barrett BA (2011) Performance, nitrogen utilization, and grazing behaviour from late-lactation dairy cows offered a fresh allocation of a ryegrass-based pasture either in the morning or in the afternoon. In ‘Adding to the knowledge base for the nutrient manager’. Occasional Report No. 24. (Eds LD Currie, CL Christensen) p. 12 (Fertilizer and Lime Research Centre, Massey University: Palmerston North, New Zealand)

Wade MH, Eirin M, Gregorini P (2006) The effect of fasting on the feeding station behavior of grazing beef heifers. Journal of Animal Science 84, 51 [Abstr.]

Woods SC, Strubbe JH (1994) The psychobiology of meals. Psychonomic Bulletin & Review 1, 141–155.
The psychobiology of meals.Crossref | GoogleScholarGoogle Scholar |

Wright W, Illius AW (1995) A comparative study of the fracture properties of five grasses. Functional Ecology 9, 269–278.
A comparative study of the fracture properties of five grasses.Crossref | GoogleScholarGoogle Scholar |

Wurtman RJ, Axelrod J (1965) The Pineal gland. Scientific American 213, 50–60.
The Pineal gland.Crossref | GoogleScholarGoogle Scholar | 1:STN:280:DyaF2M7gtlentw%3D%3D&md5=a1e212ed2f297d5d92d17e4af7dfa517CAS |

Wurtman RJ, Wurtman JJ (1989) Carbohydrates and depression. Scientific American 260, 68–75.
Carbohydrates and depression.Crossref | GoogleScholarGoogle Scholar | 1:STN:280:DyaL1M%2FosVGluw%3D%3D&md5=2b75cc725721aee48f5d0f31fcd3d954CAS |