It has been shown that cerebral lateralization is a widespread phenomenon in the
animal kingdom. Functional and structural differences between left and right brain hemispheres can be found in many other vertebrates and also in invertebrates. It has been proposed that negative, withdrawal-associated emotions are processed predominantly by the right hemisphere, whereas the left hemisphere is largely responsible for processing positive, approach-related emotions. This has been called the "laterality-
valence hypothesis". One sub-set of laterality in animals is limb dominance. Preferential limb use for specific tasks has been shown in species including chimpanzees, mice, bats, wallabies, parrots, chickens and toads. due to which, in any population, half of the mice become left-handed while the other half becomes right-handed. The learning occurs by a gradual reinforcement of randomly occurring weak asymmetries in paw choice early in training, even when training in an unbiased world. Meanwhile, reinforcement relies on short-term and long-term memory skills that are strain-dependent, Regardless of the amount of past training and consequent biasing of paw choice, there is a degree of randomness in paw choice that is not removed by training, which may provide adaptability to changing environments.
In other mammals Domestic horses (
Equus caballus) exhibit laterality in at least two areas of neural organization, i.e. sensory and motor. In
thoroughbreds, the strength of motor laterality increases with age. Horses under 4 years old have a preference to initially use the right nostril during olfaction. Along with olfaction, French horses have an eye laterality when looking at novel objects. There is a correlation between their score on an emotional index and eye preference; horses with higher emotionality are more likely to look with their left eye. The less emotive French saddlebreds glance at novel objects using the right eye, however, this tendency is absent in the
trotters, although the emotive index is the same for both breeds.
Racehorses exhibit laterality in stride patterns as well. They use their preferred stride pattern at all times whether racing or not, unless they are forced to change it while turning, injured, or fatigued. Fearfulness is an undesirable trait in guide dogs, therefore, testing for laterality can be a useful predictor of a successful guide dog. Knowing a guide dog's laterality can also be useful for training because the dog may be better at walking to the left or the right of their blind owner.
Domestic cats (
Felis catus) show an individual handedness when reaching for static food. In one study, 46% preferred to use the right paw, 44% the left, and 10% were ambi-lateral; 60% used one paw 100% of the time. There was no difference between male and female cats in the proportions of left and right paw preferences. In moving-target reaching tests, cats have a left-sided behavioural asymmetry. One study indicates that laterality in this species is strongly related to temperament. Furthermore, individuals with stronger paw preferences are rated as more confident, affectionate, active, and friendly.
Chimpanzees show right-handedness in certain conditions. This is expressed at the population level for females, but not males. The complexity of the task has a dominant effect on handedness in chimps.
Cattle use
visual/brain lateralisation in their visual scanning of novel and familiar stimuli. Domestic cattle prefer to view novel stimuli with the left eye, (similar to horses, Australian magpies, chicks, toads and fish) but use the right eye for viewing familiar stimuli.
Schreibers' long-fingered bat is lateralized at the population level and shows a left-hand bias for climbing or grasping.
In marsupials Marsupials are fundamentally different from other mammals in that they lack a
corpus callosum. However, wild
kangaroos and other
macropod marsupials have a left-hand preference for everyday tasks. Left-handedness is particularly apparent in the
red kangaroo (
Macropus rufus) and the
eastern gray kangaroo (
Macropus giganteus). The
red-necked wallaby (
Macropus rufogriseus) preferentially uses the left hand for behaviours that involve fine manipulation, but the right for behaviours that require more physical strength. There is less evidence for handedness in
arboreal species.
In birds Parrots tend to favor one foot when grasping objects (for example fruit when feeding). Some studies indicate that most parrots are left footed. The
Australian magpie (
Gymnorhina tibicen) uses both left-eye and right-eye laterality when performing anti-predator responses, which include
mobbing. Prior to withdrawing from a potential predator, Australian magpies view the animal with the left eye (85%), but prior to approaching, the right eye is used (72%). The left eye is used prior to jumping (73%) and prior to circling (65%) the predator, as well as during circling (58%) and for high alert inspection of the predator (72%). The researchers commented that "mobbing and perhaps circling are agonistic responses controlled by the LE[left eye]/right hemisphere, as also seen in other species. Alert inspection involves detailed examination of the predator and likely high levels of fear, known to be right hemisphere function."
Yellow-legged gull (
Larus michahellis) chicks show laterality when reverting from a supine to prone posture, and also in pecking at a dummy parental bill to beg for food. Lateralization occurs at both the population and individual level in the reverting response and at the individual level in begging. Females have a leftward preference in the righting response, indicating this is sex dependent. Laterality in the begging response in chicks varies according to laying order and matches variation in egg
androgens concentration.
In fish Laterality determines the organisation of
rainbowfish (
Melanotaenia spp.) schools. These fish demonstrate an individual eye preference when examining their reflection in a mirror. Fish which show a right-eye preference in the mirror test prefer to be on the left side of the school. Conversely, fish that show a left-eye preference in the mirror test or were non-lateralised, prefer to be slightly to the right side of the school. The behaviour depends on the species and sex of the school.
In amphibians Three species of toads, the
common toad (
Bufo bufo),
green toad (
Bufo viridis) and the
cane toad (
Bufo marinus) show stronger escape and defensive responses when a model predator was placed on the toad's left side compared to their right side.
Emei music frogs (
Babina daunchina) have a right-ear preference for positive or neutral signals such as a conspecific's advertisement call and white noise, but a left-ear preference for negative signals such as predatory attack.
In invertebrates The
Mediterranean fruit fly (
Ceratitis capitata) exhibits left-biased population-level lateralisation of aggressive displays (boxing with forelegs and wing strikes) with no sex-differences. In ants,
Temnothorax albipennis (rock ant) scouts show behavioural lateralization when exploring unknown nest sites, showing a population-level bias to prefer left turns. One possible reason for this is that its environment is partly maze-like and consistently turning in one direction is a good way to search and exit mazes without getting lost. This turning bias is correlated with slight asymmetries in the ants' compound eyes (differential ommatidia count). ==See also==