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Krishnan, A., Gandour, J. T., Ananthakrishnan, S., Bidelman, G. M., & Smalt, C. J. (). Functional ear (a)symmetry in brainstem neural activity relevant to encoding of voice pitch: A precursor for hemispheric specialization? Brain and Language, In Press, Corrected Proof.
Abstract: Pitch processing is lateralized to the right hemisphere; linguistic pitch is further mediated by left cortical areas. This experiment investigates whether ear asymmetries vary in brainstem representation of pitch depending on linguistic status. Brainstem frequency-following responses (FFRs) were elicited by monaural stimulation of the left and right ear of 15 native speakers of Mandarin Chinese using two synthetic speech stimuli that differ in linguistic status of tone. One represented a native lexical tone (Tone 2: T2); the other, T2', a nonnative variant in which the pitch contour was a mirror image of T2 with the same starting and ending frequencies. Two 40-ms portions of f0 contours were selected in order to compare two regions (R1, early; R2 late) differing in pitch acceleration rate and perceptual saliency. In R2, linguistic status effects revealed that T2 exhibited a larger degree of FFR rightward ear asymmetry as reflected in f0 amplitude relative to T2'. Relative to midline (ear asymmetry = 0), the only ear asymmetry reaching significance was that favoring left ear stimulation elicited by T2'. By left- and right-ear stimulation separately, FFRs elicited by T2 were larger than T2' in the right ear only. Within T2', FFRs elicited by the earlier region were larger than the later in both ears. Within T2, no significant differences in FFRS were observed between regions in either ear. Collectively, these findings support the idea that origins of cortical processing preferences for perceptually-salient portions of pitch are rooted in early, preattentive stages of processing in the brainstem.
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Lemasson, A., Koda, H., Kato, A., Oyakawa, C., Blois-Heulin, C., & Masataka, N. (2010). Influence of sound specificity and familiarity on Japanese macaques' (Macaca fuscata) auditory laterality. Behav. Brain. Res., 208(1), 286–289.
Abstract: Despite attempts to generalise the left hemisphere-speech association of humans to animal communication, the debate remains open. More studies on primates are needed to explore the potential effects of sound specificity and familiarity. Familiar and non-familiar nonhuman primate contact calls, bird calls and non-biological sounds were broadcast to Japanese macaques. Macaques turned their heads preferentially towards the left (right hemisphere) when hearing conspecific or familiar primates supporting hemispheric specialisation. Our results support the role of experience in brain organisation and the importance of social factors to understand laterality evolution.
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Carter, C., & Greening, L. Auditory stimulation of the stabled equine; the effect of different music genres on behaviour.
Abstract: Having a radio playing during the daytime at some equine establishments is common practice, but few studies have investigated whether particular music genres could be enriching for stabled horses or whether they may be perceived as aversive. This study aimed to establish whether behavioural responses differed when exposed to musical genres (Classical, Country, Jazz and Rock) and when compared to a control (no music). Eight Thoroughbred geldings (age range 8-10 years, average 8.9 years) were exposed to four musical environments and the control environment (no music) and observed in their usual stable, using instantaneous focal sampling every thirty seconds according to a pre-determined ethogram. Each horse was exposed to each genre for an hour in total, at a time when there was no human traffic or interference on the yard. All horses had been stabled for three hours before the study began. The association between genres and behavioural frequencies recorded for each environment was tested using Fisher’s Exact test of association (P<0.01), IBM SPSS21. No statistically significant associations (P=1.0) were recorded between alert or relax behaviours in Country, Classical, and Control environments. Significant associations (P<0.001) between frequency of alert behaviours and Jazz and Rock environments were noted. The latter genres appeared the most aversive which may be due to fast tempo and minor key, especially in the Jazz piece used. Country and Classical genres were slow tempo with a major key and appeared to result in more restful behaviours than Jazz or Rock. Further research is needed to; assess whether music could be used as an enriching tool, and investigate equine emotional capabilities to understand the emotional effects of music. Future studies could also consider how music impacts upon the behaviour of the human handler and whether this influences equine behaviour.
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Heffner, R. S., & Heffner, H. E. (1986). Localization of tones by horses: use of binaural cues and the role of the superior olivary complex. Behav Neurosci, 100(1), 93–103.
Abstract: The ability of horses to use binaural time and intensity difference cues to localize sound was assessed in free-field localization tests by using pure tones. The animals were required to discriminate the locus of a single tone pip ranging in frequency from 250 Hz to 25 kHz emitted by loudspeakers located 30 degrees to the left and right of the animals' midline (60 degrees total separation). Three animals were tested with a two-choice procedure; 2 additional animals were tested with a conditioned avoidance procedure. All 5 animals were able to localize 250 Hz, 500 Hz, and 1 kHz but were completely unable to localize 2 kHz and above. Because the frequency of ambiguity for the binaural phase cue delta phi for horses in this test was calculated to be 1.5 kHz, these results indicate that horses can use binaural time differences in the form of delta phi but are unable to use binaural intensity differences. This finding was supported by an unconditioned orientation test involving 4 additional horses, which showed that horses correctly orient to a 500-Hz tone pip but not to an 8-kHz tone pip. Analysis of the superior olivary complex, the brain stem nucleus at which binaural interactions first take place, reveals that the lateral superior olive (LSO) is relatively small in the horse and lacks the laminar arrangement of bipolar cells characteristic of the LSO of most mammals that can use binaural delta I.
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Harland, M. M., Stewart, A. J., Marshall, A. E., & Belknap, E. B. (2006). Diagnosis of deafness in a horse by brainstem auditory evoked potential. Can Vet J, 47(2), 151–154.
Abstract: Deafness was confirmed in a blue-eyed, 3-year-old, overo paint horse by brainstem auditory evoked potential. Congenital inherited deafness associated with lack of facial pigmentation was suspected. Assessment of hearing should be considered, especially in paint horses, at the time of pre-purchase examination. Brainstem auditory evoked potential assessment is well tolerated and accurate.
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Friederici, A. D., & Alter, K. (2004). Lateralization of auditory language functions: a dynamic dual pathway model. Brain Lang, 89(2), 267–276.
Abstract: Spoken language comprehension requires the coordination of different subprocesses in time. After the initial acoustic analysis the system has to extract segmental information such as phonemes, syntactic elements and lexical-semantic elements as well as suprasegmental information such as accentuation and intonational phrases, i.e., prosody. According to the dynamic dual pathway model of auditory language comprehension syntactic and semantic information are primarily processed in a left hemispheric temporo-frontal pathway including separate circuits for syntactic and semantic information whereas sentence level prosody is processed in a right hemispheric temporo-frontal pathway. The relative lateralization of these functions occurs as a result of stimulus properties and processing demands. The observed interaction between syntactic and prosodic information during auditory sentence comprehension is attributed to dynamic interactions between the two hemispheres.
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Lampe, J. F., & Andre, J. (2012). Cross-modal recognition of human individuals in domestic horses (Equus caballus). Animal Cognition, 15(4), 623–630.
Abstract: This study has shown that domestic horses are capable of cross-modal recognition of familiar humans. It was demonstrated that horses are able to discriminate between the voices of a familiar and an unfamiliar human without seeing or smelling them at the same moment. Conversely, they were able to discriminate the same persons when only exposed to their visual and olfactory cues, without being stimulated by their voices. A cross-modal expectancy violation setup was employed; subjects were exposed both to trials with incongruent auditory and visual/olfactory identity cues and trials with congruent cues. It was found that subjects responded more quickly, longer and more often in incongruent trials, exhibiting heightened interest in unmatched cues of identity. This suggests that the equine brain is able to integrate multisensory identity cues from a familiar human into a person representation that allows the brain, when deprived of one or two senses, to maintain recognition of this person.
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Gentner, T. Q., Fenn, K. M., Margoliash, D., & Nusbaum, H. C. (2006). Recursive syntactic pattern learning by songbirds. Nature, 440(7088), 1204–1207.
Abstract: Humans regularly produce new utterances that are understood by other members of the same language community. Linguistic theories account for this ability through the use of syntactic rules (or generative grammars) that describe the acceptable structure of utterances. The recursive, hierarchical embedding of language units (for example, words or phrases within shorter sentences) that is part of the ability to construct new utterances minimally requires a 'context-free' grammar that is more complex than the 'finite-state' grammars thought sufficient to specify the structure of all non-human communication signals. Recent hypotheses make the central claim that the capacity for syntactic recursion forms the computational core of a uniquely human language faculty. Here we show that European starlings (Sturnus vulgaris) accurately recognize acoustic patterns defined by a recursive, self-embedding, context-free grammar. They are also able to classify new patterns defined by the grammar and reliably exclude agrammatical patterns. Thus, the capacity to classify sequences from recursive, centre-embedded grammars is not uniquely human. This finding opens a new range of complex syntactic processing mechanisms to physiological investigation.
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Heffner, R. S., & Heffner, H. E. (1983). Hearing in large mammals: Horses (Equus caballus) and cattle (Bos taurus). Behavioral Neuroscience, 97(2), 299–309.
Abstract: Determined behavioral audiograms for 3 horses and 2 cows. Horses' hearing ranged from 55 Hz to 33.3 kHz, with a region of best sensitivity from 1 to 16 kHz. Cattle hearing ranged from 23 Hz to 35 kHz, with a well-defined point of best sensitivity at 8 kHz. Of the 2 species, cattle proved to have more acute hearing, with a lowest threshold of –21 db (re 20 μN/m–2) compared with the horses' lowest threshold of 7 db. Comparative analysis of the hearing abilities of these 2 species with those of other mammals provides further support for the relation between interaural distance and high-frequency hearing and between high- and low-frequency hearing. (39 ref) (PsycINFO Database Record (c) 2012 APA, all rights reserved)
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Hauber, M. E., Pearson, H. E., Reh, A., & Merges, A. (2002). Discrimination between host songs by brood parasitic brown-headed cowbirds ( Molothrus ater). Anim. Cogn., 5(3), 129–137.
Abstract: Songbirds can learn both to produce and to discriminate between different classes of acoustic stimuli. Varying levels of auditory discrimination may improve the fitness of individuals in certain ecological and social contexts and, thus, selection is expected to mold the cognitive abilities of different species according to the potential benefits of acoustic processing. Although fine-scale auditory discrimination of conspecific songs and calls has been frequently reported for brood parasitic brown-headed cowbirds ( Molothrus ater), it remains unclear why and how they perceive differently the songs of their many host species. Using habituation-dishabituation paradigms and measuring behavioral and physiological (heart-rate) responses, we found that captive female cowbirds consistently discriminated between songs of two host species, the song sparrow ( Melospiza melodia) and the red-winged blackbird ( Agelaius phoeniceus). Playback experiments with stimuli composed of con-specific followed by heterospecific vocalizations in the field also demonstrated discrimination between these heterospecific songs even though cowbirds were not attracted to playbacks of either host species' songs alone. Our results do not directly support a nest-searching function of heterospecific song discrimination by cowbirds and are most consistent with a function of the parasites' avoidance of attacks by their aggressive hosts. These data demonstrate discrimination between heterospecific vocalizations by brown-headed cowbirds and add a novel dimension to the already expansive auditory perceptual abilities of brood parasitic species and other songbirds.
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