What can whiskers tell us about mammalian evolution, behaviour, and ecology?

Most mammals have whiskers; however, nearly everything we know about whiskers derives from just a handful of species, including laboratory rats Rattus norvegicus and mice Mus musculus, as well as some species of pinniped and marsupial. We explore the extent to which the knowledge of the whisker system from a handful of species applies to mammals generally. This will help us understand whisker evolution and function, in order to gain more insights into mammalian behaviour and ecology. This review is structured around Tinbergen’s four questions, since this method is an established, comprehensive, and logical approach to studying behaviour. We ask: how do whiskers work, develop, and evolve? And what are they for? While whiskers are all slender, curved, tapered, keratinised hairs that transmit vibrotactile information, we show that there are marked differences between species with respect to whisker arrangement, numbers, length, musculature, development, and growth cycles. The conservation of form and a common muscle architecture in mammals suggests that early mammals had whiskers. Whiskers may have been functional even in therapsids. However, certain extant mammalian species are equipped with especially long and sensitive whiskers, in particular nocturnal, arboreal species, and aquatic species, which live in complex environments and hunt moving prey. Knowledge of whiskers and whisker use can guide us in developing conservation protocols and designing enriched enclosures for captive mammals. We suggest that further comparative studies, embracing a wider variety of mammalian species, are required before one can make large-scale predictions relating to evolution and function of whiskers. More research is needed to develop robust techniques to enhance the welfare and conservation of mammals

Richard Freeman Mark 1934-2003

Richard Freeman Mark was born in New Zealand and studied Medicine at Otago University, followed by doctoral studies at the Université d'Aix-Marseille in France. He undertook postdoctoral studies at the Californian Institute of Technology before accepting a Senior Lectureship at Monash University, Melbourne. His research interests focused on neuroscience, with cutting-edge studies on memory, nerve regeneration, neurodevelopment and plasticity. Richard was appointed to the Foundation Chair of Behavioural Biology at the Australian National University in 1975 and remained there for over twenty-five years. He championed an interdisciplinary and integrated approach to neurobiology in both teaching and research. He was a gifted supervisor and teacher and and initiated the first honours Neuroscience course in Australia. He was elected to the Fellowship of the Australian Academy of Science in 1974, served as President of the Australian Neuroscience Society from 1998-1999 and was awarded the Centenary Medal in 2003

Correlations Between Hand Preference and Cortical Thickness in the Secondary Somatosensory (SII) Cortex of the Common Marmoset, Callithrix jacchus

Cortical asymmetries are well established in humans for language and motor regions and correlate with handedness. Here the authors investigate structural differences in the hemispheres of left- and right-handed common marmosets using surface photography and histology. The hand preferences of 11 marmosets were assessed over their adult life span using a simple reaching task. A significant correlation was found between the length of the right lateral sulcus/brain weight and the % right-hand preference (r = .86, p = .001). Cortical thickness on the superior bank of the right lateral sulcus posteriorly was also positively correlated with % right-hand preference (r = .69, p = .025). Comparison of this site with previously published functional maps of the marmoset cortex show this area corresponds to SII, a region involved in tactile processing and somatosensory discriminations. It is suggested that the correlation between SII thickness and right-hand preference would be consistent with the fact that right-handed marmosets are more proactive than left-handers in exploring novel objects by touch. Enlargement of a cortical area involved tactile discriminations could be a precursor to the evolution of right-handedness as a population bias. © 2008 American Psychological Association

Crash characteristics of older pedestrian fatalities: Dementia pathology may be related to 'at risk' traffic situations

Older people are over represented among pedestrian casualties, and cognitive decline is an often cited possible contributory factor. Cognitive decline and dementia are intimately associated, however the role dementia might play in older pedestrian crashes has received little attention. This study describes crash characteristics for 52 fatally injured older pedestrians in the Sydney metropolitan area. It investigates the relationship between the extent of neurofibrillary tangles (NFT), a hallmark of Alzheimer's disease in the brain, and particular crash situations. The results demonstrate crash characteristics that are similar to that reported in other studies of older pedestrians. Furthermore, the results suggest that cognitive decline associated with dementia related neuropathology may be associated with specific crash situations. Compared to older pedestrians with no, or low NFT, those with moderate to high NFT were more likely to be: at least partially responsible for the incident; injured while in low complexity situations; involved in impacts with reversing vehicles; impacted in near lanes of traffic; and struck by a vehicle off road. While described as trends only (p < 0.2), these findings highlight areas of concern for older pedestrians and suggest potential targets for engineering and behaviour-based countermeasures aimed at reducing casualty numbers among older pedestrians. © 2007 Elsevier Ltd. All rights reserved

Whisker maps in marsupials: Nerve lesions and critical periods

In the wallaby, whisker-related patterns develop over a protracted period of postnatal maturation in the pouch. Afferents arrive simultaneously in the thalamus and cortex from postnatal day (P) 15. Whisker-related patterns are first seen in the thalamus at P50 and are well formed by P73, before cortical patterns first appear (P75) or are well developed (P85). This study used the slow developmental sequence and accessibility of the pouch young to investigate the effect of nerve lesions before afferent arrival, or at times when thalamic patterns are obvious but cortical patterns not yet formed. The left infraorbital nerve supplying the whiskers was cut at P0-93 and animals were perfused at P112-123. Sections through the thalamus (horizontal plane) and cortex (tangential) were reacted for cytochrome oxidase to visualize whisker-related patterns. Lesions of the nerve at P2-5, before innervation of the thalamus or cortex, resulted in an absence of patterns at both levels. Lesions from P66-77 also disrupted thalamic and cortical patterns, despite the fact that thalamic patterns are normally well established by P73. Lesions from P82-93 resulted in normal thalamic and cortical patterns. Thus, despite the wallaby having clearly separated times for the development of patterns at different levels of the pathway, these results suggest a single critical period for the thalamus and cortex, coincident with the maturation of the cortical pattern. Possible mechanisms underpinning this critical period could include dependence of the thalamic pattern on corticothalamic activity or peripheral signals to allow consolidation of thalamic barreloids. © 2006 Wiley-Liss, Inc

Electrical perceptual threshold testing: A validation study

Background/Objective: To investigate inter-rater and intra-rater reliability of electrical perceptual threshold (EPT) testing in assessing somatosensory function in healthy volunteers. Study Design: Prospective experimental. Setting: Hospital-based spinal cord injuries unit. Methods: Cutaneous electrical stimulation of 4 dermatomes at ASIA sensory key points (C3, T1, L3, and S2) was performed on 40 control subjects. The lowest ascending stimulus intensity at which sensation was perceived was recorded as the EPT. Mean EPT values for each dermatome, as determined by 2 testers at 2 time points, were examined and plotted against a normative template. Differences and associations between intra- and inter-rater measurements and left-right measurements were studied. EPT results for 2 people with spinal cord injuries were also examined. Results: EPT measurements from left and right sides, obtained from the 2 time points and 2 testers, were found to be strongly associated, with the exception of left and right side measurements at the S2 dermatome. No significant differences in the mean EPT for tester or time period were found. The intra- and inter-rater reliability was good for all dermatomes tested. Mean EPT measurements fell within the range of a normative template at each of the 4 dermatomes tested. Conclusion: EPT is an objective, reproducible, and quantifiable method of assessing sensation in a control group. However, caution should be applied in certain dermatomes such as S2, where there was large variation between left and right side measurements. © 2009 by the American Paraplegia Society

Increased neurofibrillary tangles in the brains of older pedestrians killed in traffic accidents

Background/Aims: Older people are over-represented in pedestrian fatalities, and it has been suggested that the presence of cognitive impairment or dementia in these individuals may contribute to their accidents. Using neuropathological methods, we aimed to compare the prevalence of dementia pathology in fatally injured older pedestrians with similarly aged ambulatory subjects who died from other causes. Met hods: The brains of 52 pedestrians (65-93 years) and 52 controls (65-92 years) were assessed for neurofibrillary tangles (NFT), neuritic plaques, Lewy bodies and vascular lesions using established neuropathological criteria. Results: The examination for Alzheimer's disease (AD) pathology showed that 43% of the pedestrians had NFT scores of III-VI using Braak and Braak staging, com pared with 23% of the controls (p < 0.05, Fisher's exact test), indicating incipient, possible or probable AD. There were no differences in the prevalence of pathology for vascular dementia or dementia with Lewy bodies. Conclusion:These results suggest that cognitive decline associated with AD, even in the earliest stages of the disease, may be a factor in fatal traffic accidents for older pedestrians. Special measures for pedestrian safety are necessary in areas with high densities of older citizens and especially for those diagnosed as having a mild cognitive impairment or AD. Copyright © 2006 S. Karger AG

Mild neuritic changes are increased in the brains of fatally injured older motor vehicle drivers

Given the expected increase in the older population and driving in this age group, concerns have been raised about the safety of older drivers. People over 65 years are over-represented in motor vehicle fatalities when calculated by distance driven. They are also at risk of neurodegenerative diseases, such as Alzheimer's disease, that affect cognitive function. We have examined the brains of older drivers (15M:12F) who died as a result of a motor vehicle accident (MVA) to determine the extent of Alzheimer's disease-related neurofibrillary changes (neuritic plaques and neurofibrillary tangles), Lewy body pathology and cerebrovascular disease and compared them to a control group of older licenced drivers (23M:5F) who died of other causes. The prevalence of moderate or severe neuritic plaque pathology was less than expected for the general population of this age and there was no difference between the groups. However, mild neuritic plaque pathology was increased for MVA deaths compared to controls. There was no evidence of vascular dementia or dementia with Lewy bodies. The current mandatory age-related re-licencing procedures in NSW may contribute to the low percentage of drivers with severe pathology. Further research into the role of mild pathology in cognitive impairment and older drivers is warranted. © 2007 Elsevier Ltd. All rights reserved

Delayed olfactory ensheathing cell transplants reduce nociception after dorsal root injury

Injury to cervical dorsal roots mimics the deafferentation component of brachial plexus injury in humans, with intractable neuropathic pain in the deafferented limb being a common consequence. Such lesions are generally not amenable to surgical repair. The use of olfactory ensheathing cells (OECs) for dorsal root repair, via acute transplantation, has been successful in several studies. From a clinical point of view, delayed transplantation of OECs would provide a more realistic timeframe for repair. In this study we investigated the effect of delayed OEC transplantation on functional recovery of skilled forepaw movements and amelioration of neuropathic pain, using a C7 and C8 dorsal root injury rat model previously established in our lab. We found that OEC transplantation to the dorsal horn 1. week after root injury effectively attenuated neuropathic disturbances associated with dorsal root injury, including spontaneous pain behavior, tactile allodynia and thermal hyperalgesia. The sensory controls of complex, goal-oriented skilled reaching and ladder walking, however, were not improved by delayed OEC transplantation. We did not detect any significant influence of transplanted OECs on injury-induced central reorganisation and afferent sprouting. The anti-nociceptive effect mediated by OEC transplants may therefore be explained by alternative mechanisms such as modification of inflammation and astrogliosis. The significant effect of OEC transplants in mitigating neuropathic pain may be clinically useful in intractable pain syndromes arising from deafferentation. This article is part of a Special Issue entitled: Understanding olfactory ensheathing glia and their prospect for nervous system repair. © 2010