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	Author	Dreier, S.; van Zweden, J.S.; D'Ettorre, P.
	Title	Long-term memory of individual identity in ant queens			Type	Journal Article
	Year	2007	Publication	Biology Letters	Abbreviated Journal	Biol Lett
	Volume	3	Issue	5	Pages	459-462
	Keywords	Aggression; Animals; Ants/physiology; Conditioning, Operant; Evolution; Female; Memory; *Recognition (Psychology); Social Dominance
	Abstract	Remembering individual identities is part of our own everyday social life. Surprisingly, this ability has recently been shown in two social insects. While paper wasps recognize each other individually through their facial markings, the ant, Pachycondyla villosa, uses chemical cues. In both species, individual recognition is adaptive since it facilitates the maintenance of stable dominance hierarchies among individuals, and thus reduces the cost of conflict within these small societies. Here, we investigated individual recognition in Pachycondyla ants by quantifying the level of aggression between pairs of familiar or unfamiliar queens over time. We show that unrelated founding queens of P. villosa and Pachycondyla inversa store information on the individual identity of other queens and can retrieve it from memory after 24h of separation. Thus, we have documented for the first time that long-term memory of individual identity is present and functional in ants. This novel finding represents an advance in our understanding of the mechanism determining the evolution of cooperation among unrelated individuals.
	Address	Institute of Biology, Department of Population Biology, University of Copenhagen, Universitetsparken 15, 2100 Copenhagen, Denmark. sdreier@bi.ku.dk
	Corporate Author				Thesis
	Publisher		Place of Publication		Editor
	Language	English	Summary Language		Original Title
	Series Editor		Series Title		Abbreviated Series Title
	Series Volume		Series Issue		Edition
	ISSN	1744-9561	ISBN		Medium
	Area		Expedition		Conference
	Notes	PMID:17594958			Approved	no
	Call Number	Equine Behaviour @ team @			Serial	4649
Permanent link to this record



	Author	Blackmore, T.L.; Foster, T.M.; Sumpter, C.E.; Temple, W.
	Title	An investigation of colour discrimination with horses (Equus caballus)			Type	Journal Article
	Year	2008	Publication	Behavioural Processes	Abbreviated Journal	Behav. Process.
	Volume	78	Issue	3	Pages	387-396
	Keywords	Chromatic discrimination; Colour vision; Horse; Operant
	Abstract	The ability of four horses (Equus caballus) to discriminate coloured (three shades of blue, green, red, and yellow) from grey (neutral density) stimuli, produced by back projected lighting filters, was investigated in a two response forced-choice procedure. Pushes of the lever in front of a coloured screen were occasionally reinforced, pushes of the lever in front of a grey screen were never reinforced. Each colour shade was randomly paired with a grey that was brighter, one that was dimmer, and one that approximately matched the colour in terms of brightness. Each horse experienced the colours in a different order, a new colour was started after 85% correct responses over five consecutive sessions or if accuracy showed no trend over sessions. All horses reached the 85% correct with blue versus grey, three horses did so with both yellow and green versus grey. All were above chance with red versus grey but none reached criterion. Further analysis showed the wavelengths of the green stimuli used overlapped with the yellow. The results are consistent with histological and behavioural studies that suggest that horses are dichromatic. They differ from some earlier data in that they indicate horses can discriminate yellow and blue, but that they may have deficiencies in discriminating red and green.
	Address
	Corporate Author				Thesis
	Publisher		Place of Publication		Editor
	Language		Summary Language		Original Title
	Series Editor		Series Title		Abbreviated Series Title
	Series Volume		Series Issue		Edition
	ISSN	0376-6357	ISBN		Medium
	Area		Expedition		Conference
	Notes				Approved	no
	Call Number	Equine Behaviour @ team @			Serial	5336
Permanent link to this record



	Author	Lee, J.; Floyd, T.; Erb, H.; Houpt, K.
	Title	Preference and demand for exercise in stabled horses			Type	Journal Article
	Year	2011	Publication	Applied Animal Behaviour Science	Abbreviated Journal	Appl. Anim. Behav. Sci.
	Volume	130	Issue	3-4	Pages	91-100
	Keywords	Horse; Welfare; Exercise; Operant conditioning; Two choice preference; Treadmill
	Abstract	Operant conditioning and two choice preference tests were used to assess the motivation of horses to be released from straight and from box stalls. The motivations for food, a companion, and release into a paddock were compared when the horses had to work for each commodity at increasing fixed ratios of responses (panel presses) to reward in an equine operant conditioning stall. The motivation for food (mean ± SEM = 258 ± 143) responses was much greater than that for either release (38 ± 32) from a straight stall into a large paddock alone or into a small paddock with another horse (95 ± 41) (P = 0.04). When given a two choice preference test between exercise on a treadmill for 20 min or returning to their box stalls, eight of nine horses chose to return to their stalls. In a two choice preference test six of eight horses in box stalls chose to be released into a paddock alone. Horses were given a series of two choice preference tests to determine how long they preferred to be in a paddock. After 15 min in the paddock the horses were re-tested, but all chose the paddock when released into a paddock with three other horses. They were retested every 15 min until they chose to return to their stalls. They chose to stay out for 35 ± 6 min when other horses were in the paddock but for only 17 ± 2 min when they would be alone. When deprived of stall release for 48 h the horses chose to remain in the paddock with other horses for 54 ± 6 min, but showed no compensatory behavior when they were alone (duration chosen = 16 ± 4 min). These findings indicate that horses are not strongly motivated to exercise alone and will choose not to endure forced exercise on a treadmill. The social context of voluntary exercise is important; horses are willing to stay out of their stalls longer if other horses are present and will show compensatory behavior only if other horses are present. These finding have implications for optimizing turnout time for stalled horses.
	Address
	Corporate Author				Thesis
	Publisher		Place of Publication		Editor
	Language		Summary Language		Original Title
	Series Editor		Series Title		Abbreviated Series Title
	Series Volume		Series Issue		Edition
	ISSN	0168-1591	ISBN		Medium
	Area		Expedition		Conference
	Notes				Approved	no
	Call Number	Equine Behaviour @ team @			Serial	5330
Permanent link to this record



	Author	Søndergaard, E.; Jensen, M.B.; Nicol, C.J.
	Title	Motivation for social contact in horses measured by operant conditioning			Type	Journal Article
	Year	2011	Publication	Applied Animal Behaviour Science	Abbreviated Journal	Appl. Anim. Behav. Sci.
	Volume	132	Issue	3-4	Pages	131-137
	Keywords	Horse; Housing; Social behaviour; Operant conditioning; Motivation
	Abstract	Although horses are social animals they are often housed individually with limited social contact to other horses and this may compromise their welfare. The present study included eight young female horses and investigated the strength of motivation for access to full social contact, head contact and muzzle contact, respectively, to a familiar companion horse. Horses were housed individually next to their companion horse and separations between pens prevented physical contact. During daily test sessions horses were brought to a test area where they could access an arena allowing social contact. Arena access during 3 min was given after completion of a predetermined number of responses on a panel. Fixed ratios (FR) of 8, 16, 24, 32 and 40 responses per arena access were applied in a random order, one per daily test session, within each test week (Monday to Friday), and the number of rewards per daily test session was recorded. All horses could access all three types of social contact in a cross-over design, and an empty arena was used as control. Motivational strength was assessed using elasticity of demand functions, which were estimated based on the number of rewards earned and FR. Elasticities of demand for the three types of social contact were low (-0.20), and not significantly different, although increasing FR still resulted in a decrease in rewards obtained for all three types of social contact (P < 0.001). Across FR-levels horses earned more rewards for social contact than for an empty arena, as shown by much higher intercept values (2.51 vs. 0.99; P < 0.001). However, the elasticity of demand for infrequent access to an empty arena (-0.08) was lower than for social contact (P < 0.01) and not significantly different from zero (P = 0.07). Horses performed more social behaviour the lesser the restriction on social contact (full > head > muzzle). However, the finding that horses showed a similar and high motivation for all three types of social contact suggests that they are valued equally highly in a situation where the alternative is no social contact.
	Address
	Corporate Author				Thesis
	Publisher		Place of Publication		Editor
	Language		Summary Language		Original Title
	Series Editor		Series Title		Abbreviated Series Title
	Series Volume		Series Issue		Edition
	ISSN	0168-1591	ISBN		Medium
	Area		Expedition		Conference
	Notes				Approved	no
	Call Number	Equine Behaviour @ team @			Serial	5410
Permanent link to this record



	Author	Ahrendt, L.P.; Labouriau, R.; Malmkvist, J.; Nicol, C.J.; Christensen, J.W.
	Title	Development of a standard test to assess negative reinforcement learning in horses			Type	Journal Article
	Year	2015	Publication	Applied Animal Behaviour Science	Abbreviated Journal	Appl. Anim. Behav. Sci.
	Volume	169	Issue		Pages	38-42
	Keywords	Algometry; Horse behaviour; Learning performance; Operant conditioning; Pressure-release; Horse training
	Abstract	Most horses are trained by negative reinforcement. Currently, however, no standardised test for evaluating horses' negative reinforcement learning ability is available. The aim of this study was to develop an objective test to investigate negative reinforcement learning in horses. Twenty-four Icelandic horses (3 years old) were included in this study. The horses were tested in a pressure-release task on three separate days with 10, 7 and 5 trials on each side, respectively. Each trial consisted of pressure being applied on the hindquarter with an algometer. The force of the pressure was increased until the horse moved laterally away from the point of pressure. There was a significant decrease in required force over trials on the first test day (P<0.001), but not the second and third day. The intercepts on days 2 and 3 differed significantly from day 1 (P<0.001), but not each other. Significantly stronger force was required on the right side compared to the left (P<0.001), but there was no difference between first and second side tested (P=0.56). Individual performance was evaluated by median-force and the change in force over trials on the first test day. These two measures may explain different characteristics of negative reinforcement learning. In conclusion, this study presents a novel, standardised test for evaluating negative reinforcement learning ability in horses.
	Address
	Corporate Author				Thesis
	Publisher		Place of Publication		Editor
	Language		Summary Language		Original Title
	Series Editor		Series Title		Abbreviated Series Title
	Series Volume		Series Issue		Edition
	ISSN	0168-1591	ISBN		Medium
	Area		Expedition		Conference
	Notes				Approved	no
	Call Number	Equine Behaviour @ team @			Serial	6650
Permanent link to this record



	Author	Mejdell, C.M.; Buvik, T.; Jørgensen, G.H.M.; Bøe, K.E.
	Title	Horses can learn to use symbols to communicate their preferences			Type	Journal Article
	Year	2016	Publication	Applied Animal Behaviour Science	Abbreviated Journal	Appl. Anim. Behav. Sci.
	Volume	184	Issue		Pages	66-73
	Keywords	Operant conditioning; Blanket; Rug; Thermoregulation; Cognition; Clicker training
	Abstract	Abstract This paper describes a method in which horses learn to communicate by touching different neutral visual symbols, in order to tell the handler whether they want to have a blanket on or not. Horses were trained for 10–15 min per day, following a training program comprising ten steps in a strategic order. Reward based operant conditioning was used to teach horses to approach and touch a board, and to understand the meaning of three different symbols. Heat and cold challenges were performed to help learning and to check level of understanding. At certain stages, a learning criterion of correct responses for 8–14 successive trials had to be achieved before proceeding. After introducing the free choice situation, on average at training day 11, the horse could choose between a “no change” symbol and the symbol for either “blanket on” or “blanket off” depending on whether the horse already wore a blanket or not. A cut off point for performance or non-performance was set to day 14, and 23/23 horses successfully learned the task within this limit. Horses of warm-blood type needed fewer training days to reach criterion than cold-bloods (P < 0.05). Horses were then tested under differing weather conditions. Results show that choices made, i.e. the symbol touched, was not random but dependent on weather. Horses chose to stay without a blanket in nice weather, and they chose to have a blanket on when the weather was wet, windy and cold (χ2 = 36.67, P < 0.005). This indicates that horses both had an understanding of the consequence of their choice on own thermal comfort, and that they successfully had learned to communicate their preference by using the symbols. The method represents a novel tool for studying preferences in horses.
	Address
	Corporate Author				Thesis
	Publisher		Place of Publication		Editor
	Language		Summary Language		Original Title
	Series Editor		Series Title		Abbreviated Series Title
	Series Volume		Series Issue		Edition
	ISSN	0168-1591	ISBN		Medium
	Area		Expedition		Conference
	Notes				Approved	no
	Call Number	Equine Behaviour @ team @			Serial	6022
Permanent link to this record



	Author	Mejdell, C.M.; Buvik, T.; Jørgensen, G.H.M.; Bøe, K.E.
	Title	Horses can learn to use symbols to communicate their preferences			Type	Journal Article
	Year	2016	Publication	Applied Animal Behaviour Science	Abbreviated Journal	Appl. Anim. Behav. Sci.
	Volume	184	Issue		Pages	66-73
	Keywords	Operant conditioning; Blanket; Rug; Thermoregulation; Cognition; Clicker training
	Abstract	This paper describes a method in which horses learn to communicate by touching different neutral visual symbols, in order to tell the handler whether they want to have a blanket on or not. Horses were trained for 10-15min per day, following a training program comprising ten steps in a strategic order. Reward based operant conditioning was used to teach horses to approach and touch a board, and to understand the meaning of three different symbols. Heat and cold challenges were performed to help learning and to check level of understanding. At certain stages, a learning criterion of correct responses for 8-14 successive trials had to be achieved before proceeding. After introducing the free choice situation, on average at training day 11, the horse could choose between a “no change” symbol and the symbol for either “blanket on” or “blanket off” depending on whether the horse already wore a blanket or not. A cut off point for performance or non-performance was set to day 14, and 23/23 horses successfully learned the task within this limit. Horses of warm-blood type needed fewer training days to reach criterion than cold-bloods (P<0.05). Horses were then tested under differing weather conditions. Results show that choices made, i.e. the symbol touched, was not random but dependent on weather. Horses chose to stay without a blanket in nice weather, and they chose to have a blanket on when the weather was wet, windy and cold (χ2=36.67, P<0.005). This indicates that horses both had an understanding of the consequence of their choice on own thermal comfort, and that they successfully had learned to communicate their preference by using the symbols. The method represents a novel tool for studying preferences in horses.
	Address
	Corporate Author				Thesis
	Publisher		Place of Publication		Editor
	Language		Summary Language		Original Title
	Series Editor		Series Title		Abbreviated Series Title
	Series Volume		Series Issue		Edition
	ISSN	0168-1591	ISBN		Medium
	Area		Expedition		Conference
	Notes				Approved	no
	Call Number	Equine Behaviour @ team @			Serial	6617
Permanent link to this record



	Author	Mejdell, C.M.; Buvik, T.; Jørgensen, G.H.M.; Bøe, K.E.
	Title	Horses can learn to use symbols to communicate their preferences			Type	Journal Article
	Year	2016	Publication	Applied Animal Behaviour Science	Abbreviated Journal	Appl. Anim. Behav. Sci.
	Volume	184	Issue		Pages	66-73
	Keywords	Operant conditioning; Blanket; Rug; Thermoregulation; Cognition; Clicker training
	Abstract	This paper describes a method in which horses learn to communicate by touching different neutral visual symbols, in order to tell the handler whether they want to have a blanket on or not. Horses were trained for 10-15min per day, following a training program comprising ten steps in a strategic order. Reward based operant conditioning was used to teach horses to approach and touch a board, and to understand the meaning of three different symbols. Heat and cold challenges were performed to help learning and to check level of understanding. At certain stages, a learning criterion of correct responses for 8-14 successive trials had to be achieved before proceeding. After introducing the free choice situation, on average at training day 11, the horse could choose between a “no change” symbol and the symbol for either “blanket on” or “blanket off” depending on whether the horse already wore a blanket or not. A cut off point for performance or non-performance was set to day 14, and 23/23 horses successfully learned the task within this limit. Horses of warm-blood type needed fewer training days to reach criterion than cold-bloods (P<0.05). Horses were then tested under differing weather conditions. Results show that choices made, i.e. the symbol touched, was not random but dependent on weather. Horses chose to stay without a blanket in nice weather, and they chose to have a blanket on when the weather was wet, windy and cold (χ2=36.67, P<0.005). This indicates that horses both had an understanding of the consequence of their choice on own thermal comfort, and that they successfully had learned to communicate their preference by using the symbols. The method represents a novel tool for studying preferences in horses.
	Address
	Corporate Author				Thesis
	Publisher		Place of Publication		Editor
	Language		Summary Language		Original Title
	Series Editor		Series Title		Abbreviated Series Title
	Series Volume		Series Issue		Edition
	ISSN	0168-1591	ISBN		Medium
	Area		Expedition		Conference
	Notes				Approved	no
	Call Number	Equine Behaviour @ team @			Serial	6651
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