Editing Laterality (section)

== In other animals ==
It has been shown that cerebral lateralization is a widespread phenomenon in the [[animal|animal kingdom]].<ref>Rogers, Lesley J., Andrew, Richard J. (2002) Comparative Vertebrate Lateralization, Cambridge University Press</ref> Functional and structural differences between left and right brain hemispheres can be found in many other vertebrates and also in invertebrates.<ref name="Manns">{{cite journal|author1=Manns, M. |author2=Ströckens, F.|year=2014|title=Functional and structural comparison of visual lateralization in birds–similar but still different|journal=Frontiers in Psychology|volume=5|page=206|doi=10.3389/fpsyg.2014.00206|pmid=24723898|pmc=3971188|doi-access=free}}</ref>

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 (psychology)|valence]] hypothesis".<ref name="Barnard2015">{{cite journal|author=Barnard, S., Matthews, L., Messori, S., Podaliri-Vulpiani, M. and Ferri, N.|year=2015|title=Laterality as an indicator of emotional stress in ewes and lambs during a separation test|journal=Animal Cognition|volume=19|issue=1|pages=1–8|doi=10.1007/s10071-015-0928-3|pmid=26433604|s2cid=7008274}}</ref>

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.<ref name="Manns" />

Another form of laterality is hemispheric dominance for processing conspecific vocalizations, reported for chimpanzees, sea lions, dogs, zebra finches and Bengalese finches.<ref name="Manns" />

=== In mice ===
In mice (''Mus musculus''), laterality in paw usage has been shown to be a learned behavior (rather than inherited),<ref>{{Cite journal|last1=Biddle|first1=Fred G|last2=Eales|first2=Brenda A| year = 2006 |title=Hand-preference training in the mouse reveals key elements of its learning and memory process and resolves the phenotypic complexity in the behaviour|journal=Genome|language=en|volume=49|issue=6|pages=666–677|doi=10.1139/g06-026|pmid=16936846|issn=0831-2796}}</ref> 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.<ref name=":0">{{Cite journal|last1=Ribeiro|first1=Andre S.|last2=Lloyd-Price|first2=Jason|last3=Eales|first3=Brenda A.|last4=Biddle|first4=Fred G.|year = 2010|title=Dynamic Agent-Based Model of Hand-Preference Behavior Patterns in the Mouse|journal=Adaptive Behavior|language=en|volume=18|issue=2|pages=116–131|doi=10.1177/1059712309339859|s2cid=10117297|issn=1059-7123}}</ref><ref name=":1">{{Cite journal|last1=Ribeiro|first1=Andre S.|last2=Eales|first2=Brenda A.|last3=Biddle|first3=Fred G.|year = 2011|title=Learning of paw preference in mice is strain dependent, gradual and based on short-term memory of previous reaches|journal=Animal Behaviour|language=en|volume=81|issue=1|pages=249–257|doi=10.1016/j.anbehav.2010.10.014|s2cid=26136740}}</ref> Meanwhile, reinforcement relies on short-term and long-term memory skills that are strain-dependent,<ref name=":0" /><ref name=":1" /> causing strains to differ in the degree of laterality of its individuals. Long-term memory of previously gained laterality in handedness due to training is heavily diminished in mice with absent corpus callosum and reduced hippocampal commissure.<ref>{{Cite journal|last1=Ribeiro|first1=Andre S.|last2=Eales|first2=Brenda A.|last3=Biddle|first3=Fred G.|year = 2013|title=Short-term and long-term memory deficits in handedness learning in mice with absent corpus callosum and reduced hippocampal commissure|journal=Behavioural Brain Research|language=en|volume=245|pages=145–151|doi=10.1016/j.bbr.2013.02.021|pmid=23454853|s2cid=40650630}}</ref> 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,<ref>{{Cite journal|last1=Ribeiro|first1=Andre S.|last2=Eales|first2=Brenda A.|last3=Lloyd-Price|first3=Jason|last4=Biddle|first4=Fred G.|year = 2014|title=Predictability and randomness of paw choices are critical elements in the behavioural plasticity of mouse paw preference|journal=Animal Behaviour|language=en|volume=98|pages=167–176|doi=10.1016/j.anbehav.2014.10.008|s2cid=53144817}}</ref> which may provide adaptability to changing environments.

=== In other mammals ===
[[Domestic horse]]s (''Equus caballus'') exhibit laterality in at least two areas of neural organization, i.e. sensory and motor. In [[thoroughbred]]s, the strength of motor laterality increases with age. Horses under 4 years old have a preference to initially use the right nostril during olfaction.<ref>{{Cite journal|title = Motor and sensory laterality in thoroughbred horses|author1=McGreevy, P. |author2=Rogers, L.|year= 2005|journal = Applied Animal Behaviour Science|doi = 10.1016/j.applanim.2004.11.012|issue = 4|volume = 92|pages = 337–352}}</ref> 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 [[Trotter (horse)|trotters]], although the emotive index is the same for both breeds.<ref>{{Cite journal|title = Laterality of horses associated with emotionality in novel situations|author=Larose, C., Richard-Yris, M.-A., Hausberger, M. and Rogers, L.J.|year=2006|journal = Laterality: Asymmetries of Body, Brain and Cognition|doi = 10.1080/13576500600624221|pmid = 16754236|issue = 4|pages = 355–367|volume = 11|s2cid=31432670}}</ref> [[Racehorse]]s 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.<ref>{{Cite journal|title = Laterality in stride pattern preference in racehorses|author1=Williams, D.E. |author2=Norris, B.J|year=2007|journal = Animal Behaviour|doi = 10.1016/j.anbehav.2007.01.014|issue = 4|pages = 941–950|volume = 74|s2cid = 53166627}}</ref>

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.<ref name="Tomkins">{{cite journal |author= Tomkins, L.M., Thomson, P.C. and McGreevy, P.D.|year=2010|title = First-stepping Test as a measure of motor laterality in dogs (Canis familiaris) | journal = Journal of Veterinary Behavior: Clinical Applications and Research | volume = 5 | issue = 5| pages = 247–255 | doi = 10.1016/j.jveb.2010.03.001 }}</ref>

[[Domestic cat]]s (''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.<ref>{{cite journal|journal=Behavioural Processes|volume=39|issue=3|year=1997|pages=241–247|title=Paw preferences in cats (Felis silvestris catus) living in a household environment|doi=10.1016/S0376-6357(96)00758-9|author1=Pike, A.V.L. |author2=Maitland, D.P.|pmid=24897330|s2cid=26114508}}</ref> 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.<ref name="McDowell2016">{{cite journal|author=McDowell, L.J., Wells, D.L., Hepper, P.G. and Dempster, M.|year=2016|title=Lateral bias and temperament in the domestic cat (Felis Silvestris)|journal=Journal of Comparative Psychology |volume=130|issue=4|pages=313–320|doi=10.1037/com0000030|pmid=27359075}}</ref>

[[Common chimpanzee|Chimpanzee]]s 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.<ref>{{cite journal|author=Llorente, M., Riba, D., Palou, L., Carrasco, L., Mosquera, M., Colell, M. and Feliu, O.|year=2011|title=Population-level right-handedness for a coordinated bimanual task in naturalistic housed chimpanzees: replication and extension in 114 animals from Zambia and Spain|journal=American Journal of Primatology|volume=73|issue=3|pages=281–290|doi=10.1002/ajp.20895|pmid=20954250|s2cid=24054277}}</ref>

[[Cattle]] use [[Lateralization of brain function|visual/brain lateralisation]] in their visual scanning of novel and familiar stimuli.<ref>{{cite journal |author=Phillips, C.J.C., Oevermans, H., Syrett, K.L., Jespersen, A.Y. and Pearce, G.P. |year=2015 |title=Lateralization of behavior in dairy cows in response to conspecifics and novel persons |journal=Journal of Dairy Science |volume=98 |issue=4 |pages=2389–2400 |doi=10.3168/jds.2014-8648|pmid=25648820 |doi-access=free }}</ref> 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.<ref name="Robins">{{cite journal |author1=Robins, A. |author2=Phillips, C. |year=2010 |title=Lateralised visual processing in domestic cattle herds responding to novel and familiar stimuli |journal=Laterality |volume=15 |issue=5 |pages=514–534 |doi=10.1080/13576500903049324|pmid=19629847 |s2cid=13283847 }}</ref>

[[Schreibers' long-fingered bat]] is lateralized at the population level and shows a left-hand bias for climbing or grasping.<ref>{{cite journal |last1=Zucca |first1=P. |last2=Palladini |first2=A. |last3=Baciadonna |first3=L. |last4=Scaravelli |first4=D. |year=2010 |title=Handedness in the echolocating Schreiber's long-fingered bat (''Miniopterus schreibersii'') |journal=Behavioural Processes |volume=84 |issue=3 |pages=693–695 |doi=10.1016/j.beproc.2010.04.006|pmid=20399840 |s2cid=3093349 }}</ref>

====In marsupials====
[[Marsupials]] are fundamentally different from other mammals in that they lack a [[corpus callosum]].<ref name="Nowak1999">{{cite book |last=Nowak |first=Ronald M. |title=Walker's Mammals of the World |url=https://archive.org/details/walkersmammalsof0002nowa |url-access=registration |date=1999 |publisher=Johns Hopkins University Press |isbn=978-0-8018-5789-8}}</ref> However, wild [[kangaroo]]s and other [[Macropodidae|macropod]] [[marsupial]]s 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.<ref name="Sci-News">{{cite web |title=All kangaroos are lefties, scientists say |website=Sci-News.com |url=http://www.sci-news.com/biology/science-kangaroos-lefties-02929.html |date=June 18, 2015 |access-date=June 19, 2015}}</ref>

===In birds===
[[Parrot]]s tend to favor one foot when grasping objects (for example fruit when feeding). Some studies indicate that most parrots are left footed.<ref>Zeigler, H. Phillip & Hans-Joachim Bischof, eds. ''Vision, Brain, and Behavior in Birds.'' Cambridge, MA: MIT Press, 1993. 239.</ref>

The [[Australian magpie]] (''Gymnorhina tibicen'') uses both left-eye and right-eye laterality when performing anti-predator responses, which include [[Mobbing (animal behavior)|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."<ref>{{cite journal|author=Koboroff, A., Kaplan, G. and Rogers, L.J.|year=2008|title=Hemispheric specialization in Australian magpies (Gymnorhina tibicen) shown as eye preferences during response to a predator|journal=Brain Research Bulletin|volume=76|issue=3|pages=304–306|doi=10.1016/j.brainresbull.2008.02.015|pmid=18498946|s2cid=20559048|url=http://e-publications.une.edu.au/1959.11/3059|url-access=subscription}}</ref>

[[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 [[androgen]]s concentration.<ref>{{cite journal|author=Romano, M., Parolini, M., Caprioli, M., Spiezio, C., Rubolini, D. and Saino, N.|year=2015|title=Individual and population-level sex-dependent lateralization in yellow-legged gull (Larus michahellis) chicks|journal=Behavioural Processes|volume=115|pages=109–116|doi=10.1016/j.beproc.2015.03.012|pmid=25818662|s2cid=40189333}}</ref>

===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.<ref name="Bibost">{{cite journal|author1=Bibost, A-L. |author2=Brown, C.|year=2013|title=Laterality enhances schooling position in rainbowfish, Melaotaenia spp|journal=PLOS ONE|volume=8|issue=11|pages=e80907|doi=10.1371/journal.pone.0080907|pmid=24260506|pmc=3829960|bibcode=2013PLoSO...880907B|doi-access=free}}</ref>

===In amphibians===
Three species of toads, the [[common toad]] (''Bufo bufo''), [[Bufo viridis|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.<ref>{{cite journal|author=Lippolis, G., Bisazza, A., Rogers, L. J. and Vallortigara, G.|year=2002|title=Lateralisation of predator avoidance responses in three species of toads|journal=Laterality: Asymmetries of Body, Brain and Cognition|volume=7|issue=2|pages=163–183|doi=10.1080/13576500143000221|pmid=15513195|citeseerx=10.1.1.511.7850|s2cid=14978610}}</ref> [[Emei music frog]]s (''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.<ref>{{cite journal|author=Xue, F., Fang, G., Yang, P., Zhao, E., Brauth, S. E. and Tang, Y.|year=2015|title=The biological significance of acoustic stimuli determines ear preference in the music frog|journal=The Journal of Experimental Biology|volume=218|issue=5|pages=740–747|doi=10.1242/jeb.114694|pmid=25740903|doi-access=free|bibcode=2015JExpB.218..740X }}</ref>

===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.<ref>{{cite journal|author=Benelli, G. |author2=Donati, E. |author3=Romano, D. |author4=Stefanini, C. |author5=Messing, R. H. |author6=Canale, A. |year=2015|title= Lateralisation of aggressive displays in a tephritid fly|journal=The Science of Nature|volume=102|issue=1–2|pages=1–9|doi=10.1007/s00114-014-1251-6|pmid=25599665|bibcode=2015SciNa.102....1B|s2cid=17242438}}</ref> 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.<ref>{{cite journal |vauthors=Hunt ER, etal | year=2014 |title=Ants show a leftward turning bias when exploring unknown nest sites|journal=[[Biology Letters]] |volume=10 |issue=12 |pmid=25540159 |doi=10.1098/rsbl.2014.0945 |pmc=4298197 |page=20140945}}</ref> This turning bias is correlated with slight asymmetries in the ants' compound eyes (differential ommatidia count).<ref>{{cite journal |vauthors=Hunt ER, etal | year=2018 |title=Asymmetric ommatidia count and behavioural lateralization in the ant ''Temnothorax albipennis''|journal=[[Scientific Reports]] |volume=8 |issue=5825 | page=5825 |doi=10.1038/s41598-018-23652-4| pmid=29643429 | pmc=5895843 | bibcode=2018NatSR...8.5825H }}</ref>