Abstract: Behavioural innovations have become central to our thinking about how animals adjust to changing environments. It is now well established that animals vary in their ability to innovate, but understanding why remains a challenge. This is because innovations are rare, so studying innovation requires alternative experimental assays that create opportunities for animals to express their ability to invent new behaviours, or use pre-existing ones in new contexts. Problem solving of extractive foraging tasks has been put forward as a suitable experimental assay. We review the rapidly expanding literature on problem solving of extractive foraging tasks in order to better understand to what extent the processes underpinning problem solving, and the factors influencing problem solving, are in line with those predicted, and found, to underpin and influence innovation in the wild. Our aim is to determine whether problem solving can be used as an experimental proxy of innovation. We find that in most respects, problem solving is determined by the same underpinning mechanisms, and is influenced by the same factors, as those predicted to underpin, and to influence, innovation. We conclude that problem solving is a valid experimental assay for studying innovation, propose a conceptual model of problem solving in which motor diversity plays a more central role than has been considered to date, and provide recommendations for future research using problem solving to investigate innovation. This article is part of a Special Issue entitled: Cognition in the wild.

Abstract: A nonsense mutation in DMRT3 (‘Gait keeper’ mutation) has a predominant effect on gaiting ability in horses, being permissive for the ability to perform lateral gaits and having a favourable effect on speed capacity in trot. The DMRT3 mutant allele (A) has been found in high frequency in gaited breeds and breeds bred for harness racing, while other horse breeds were homozygous for the wild-type allele (C). The aim of this study was to evaluate further the effect of the DMRT3 nonsense mutation on the gait quality and speed capacity in the multigaited Icelandic horse and demonstrate how the frequencies of the A- and C- alleles have changed in the Icelandic horse population in recent decades. It was confirmed that homozygosity for the DMRT3 nonsense mutation relates to the ability to pace. It further had a favourable effect on scores in breeding field tests for the lateral gait tölt, demonstrated by better beat quality, speed capacity and suppleness. Horses with the CA genotype had on the other hand significantly higher scores for walk, trot, canter and gallop, and they performed better beat and suspension in trot and gallop. These results indicate that the AA genotype reinforces the coordination of ipsilateral legs, with the subsequent negative effect on the synchronized movement of diagonal legs compared with the CA genotype. The frequency of the A-allele has increased in recent decades with a corresponding decrease in the frequency of the C-allele. The estimated frequency of the A-allele in the Icelandic horse population in 2012 was 0.94. Selective breeding for lateral gaits in the Icelandic horse population has apparently altered the frequency of DMRT3 genotypes with a predicted loss of the C-allele in relatively few years. The results have practical implications for breeding and training of Icelandic horses and other gaited horse breeds.

Abstract: Recent results suggest that wild Northern herbivores reduce their metabolism during times of low ambient temperatures and food shortage in order to reduce their energetic needs. It is however not known if domesticated animals are also able to reduce their energy expenditure. We exposed ten Shetland pony mares to different environmental conditions (summer and winter) and to two food quantities (60 and 100% of maintenance energy requirement, respectively) during low winter temperatures to examine energetic and behavioural responses. In summer ponies showed a considerably higher field metabolic rate (FMR) (63.4±15.0 MJ d-1) compared to restrictively fed and control animals in winter (24.6±7.8 MJ d-1 and 15.0±1.1 MJ d-1, respectively). During summer conditions locomotor activity, resting heart rates and total water turnover were considerably elevated (P<0.001) compared to winter. Restrictively fed animals (N=5) compensated for the decreased energy supply by reducing their FMR by 26% compared to control animals (N=5). Furthermore, resting heart rate, body mass and body condition score were lower (29.2±2.7 beats min-1; 140±22 kg; 3.0±1.0 points) than in control animals (36.8±41 beats min-1; 165 ±31 kg; 4.4±0.7 points; P<0.05). While the observed behaviour did not change, nocturnal hypothermia was elevated. We conclude that ponies acclimatize to different climatic conditions by changing their metabolic rate, behaviour and some physiological parameters. When exposed to energy challenges, ponies, like wild herbivores, exhibited hypometabolism and nocturnal hypothermia.

Abstract: Intraspecific allometric calculations of the brain to body size relation revealed distinct differences between 127 (67; 60) ancestral wild cavies and 82 (37; 45) guinea pigs, their domesticated relatives. The dependency of both measures from one another remained the same in both animal groups but the brains of guinea pigs were by 14.22% smaller at any net body weight. Consistent with results in other species the domestication of Cavia aperea is also characterized by a decrease of brain size. Fresh tissue sizes of the five brain parts medulla oblongata, cerebellum, mesencephalon, diencephalon and telencephalon were determined for 6 cavies and 6 guinea pigs by the serial section method. Additionally the sizes of 16 endbrain structures and those of the optic tract, the lateral geniculate body and the cochlear nucleus were measured. Different decrease values resulted for all these structures concomitant with domestication as was calculated from the amount of total brain size decrease and average relative structure values in the wild as well as the domesticated brain. The size decrease of the entire telencephalon (-13.7%) was within the range of the mean overall reduction as similarly was the case for the total neocortex (-10.7%) whereas the total allocortex (-20.9%) clearly was more strongly affected. The size decrease of the olfactory bulb (-41.9%) was extreme and clearly higher than found for the secondary olfactory structures (around -11%). The primary nuclei of other sensory systems (vision, audition) were decreased to less extent (lateral geniculate: -18.1%; cochlear nucleus: -12.6%). Mass decreases of pure white matter parts were nearly twice as high in contrast to associated grey matter parts (neocortex white versus grey matter; tractus opticus versus lateral geniculate body). The relatively great decrease values found for the limbic structures hippocampus (-26.9%) and schizocortex (-25.9%) are especially notable since they are in good conformity with domestication effects in other mammalian species. The findings of this study are discussed with regard to results of similar investigations on wild and domesticated gerbils (Meriones unguiculatus), the encephalization of the wild form, the special and species-specific mode and duration of domestication and in connection with certain behavioral changes as resulted from comparative investigations in ethology, socio-biology, endocrinology and general physiology.