Organization of the spinal trigeminal nucleus in star-nosed moles.

Somatosensory inputs from the face project to multiple regions of the trigeminal nuclear complex in the brainstem. In mice and rats, three subdivisions contain visible representations of the mystacial vibrissae, the principal sensory nucleus, spinal trigeminal subnucleus interpolaris, and subnucleus caudalis. These regions are considered important for touch with high spatial acuity, active touch, and pain and temperature sensation, respectively. Like mice and rats, the star-nosed mole (Condylura cristata) is a somatosensory specialist. Given the visible star pattern in preparations of the star-nosed mole cortex and the principal sensory nucleus, we hypothesized there were star patterns in the spinal trigeminal nucleus subnuclei interpolaris and caudalis. In sections processed for cytochrome oxidase, we found star-like segmentation consisting of lightly stained septa separating darkly stained patches in subnucleus interpolaris (juvenile tissue) and subnucleus caudalis (juvenile and adult tissue). Subnucleus caudalis represented the face in a three-dimensional map, with the most anterior part of the face represented more rostrally than posterior parts of the face. Multiunit electrophysiological mapping was used to map the ipsilateral face. Ray-specific receptive fields in adults matched the CO segmentation. The mean areas of multiunit receptive fields in subnucleus interpolaris and caudalis were larger than previously mapped receptive fields in the mole's principal sensory nucleus. The proportion of tissue devoted to each ray's representation differed between the subnucleus interpolaris and the principal sensory nucleus. Our finding that different trigeminal brainstem maps can exaggerate different parts of the face could provide new insights for the roles of these different somatosensory stations.

Mechanical performance of rat, mouse and mole spring traps, and possible implications for welfare performance.

Lethal spring traps are widely used for killing small mammals in the UK. Many require government approval, based primarily on humaneness. However, mole traps and break-back traps for rats and mice are exempt; those available vary widely in price and apparent quality. The EU is considering implementing a Trapping Directive that would alter UK legislation, and a recent report advised the EU that trapping legislation should cover all trapped species and encourage improvement of traps. Mechanical trap performance is often used as an indicator of welfare impact. We examined the mechanical evidence for scope to improve the welfare standards of rat, mouse and mole spring traps. We measured mechanical performance among a range of rat, mouse and mole traps. Impact momentum values varied 6-8 fold, and clamping force values 4-5.5 fold, among traps for killing each species. There was considerable overlap in the performance of rat and mouse traps. Trap-opening angle and spring type were related to impact momentum and clamping force in traps for both species. There was no relationship between price and mechanical performance in traps for any species, except talpa mole traps. We are unable to judge the direct welfare impact of the traps tested, but rather the potential welfare threat associated with their exemption from approval. The wide variation in mechanical performance in traps for each species, overlap in performance between rat and mouse traps and increasing availability of weaker plastic rodent traps indicate considerable scope for improving the humaneness of spring traps for rats, mice and moles. We conclude that all such traps should be subject to the UK approval process. New welfare categories might improve trap standards further. Our results could also help improve rodent trap design and assist consumers in selecting more powerful traps. Many thousands of rats, mice and moles might benefit.

[Convergent development of the subcutaneous muscles in moles and golden moles].

The goal of the investigation was to clarify the ways of subcutaneous muscles development in burrowing forms of insectivores--moles (Talpidae) and golden moles (Chrysochloridae). To achieve the goal the comparative morpho-functional analysis of subcutaneous and facial musculature of six mole genera (Neurotrichus, Urotrichus, Talpa, Mogera, Scalopus, Parascalops), two genera of chrysochlorids (Chrysochloris, Eremitalpa), and some other non-specialized forms of insectivorous mammals from Erinaceidae, Tenrecidae, Soricidae and Solenodontidae was carried out. It was shown that some of m. cutaneus trunci derivatives interact intimately with facial musculature. Besides, subcutaneous muscle forms additional layers that facilitate the operation of shaking ground particles off the for, which is rather important for burrowers. Overall complication of subcutaneous musculature in moles and chrysochlorids is accompanied by convergent similarity in the development of some musculature portions. At the same time some layers of m. cutaneus trunci seem to have evolved in different, special ways.

The molecular basis of defective lens development in the Iberian mole.

BACKGROUND: Fossorial mammals face natural selection pressures that differ from those acting on surface dwelling animals, and these may lead to reduced visual system development. We have studied eye development in a species of true mole, the Iberian mole Talpa occidentalis, and present the molecular basis of abnormal lens development. This is the first embryological developmental study of the eyes of any fossorial mammal at the molecular level. RESULTS: Lens fibre differentiation is not completed in the Iberian mole. Although eye development starts normally (similar to other model species), defects are seen after closure of the lens vesicle. PAX6 is not down-regulated in developing lens fibre nuclei, as it is in other species, and there is ectopic expression of FOXE3, a putative downstream effector of PAX6, in some, but not all lens fibres. FOXE3-positive lens fibres continue to proliferate within the posterior compartment of the embryonic lens, but unlike in the mouse, no proliferation was detected anywhere in the postnatal mole lens. The undifferentiated status of the anterior epithelial cells was compromised, and most of them undergo apoptosis. Furthermore, beta-crystallin and PROX1 expression patterns are abnormal and our data suggest that genes encoding beta-crystallins are not directly regulated by PAX6, c-MAF and PROX1 in the Iberian mole, as they are in other model vertebrates. CONCLUSION: In other model vertebrates, genetic pathways controlling lens development robustly compartmentalise the lens into a simple, undifferentiated, proliferative anterior epithelium, and quiescent, anuclear, terminally differentiated posterior lens fibres. These pathways are not as robust in the mole, and lead to loss of the anterior epithelial phenotype and only partial differentiation of the lens fibres, which continue to express 'epithelial' genes. Paradigms of genetic regulatory networks developed in other vertebrates appear not to hold true for the Iberian mole.

The evolution of female mole ovotestes evidences high plasticity of mammalian gonad development.

Previous studies of the reproductive biology and genetics of European moles (Talpa spp.) showed that all females of these species have ovotestes (gonads with testicular and ovarian tissue) instead of normal ovaries, a unique specialization among mammals. Females are fertile as their ovarian tissue is fully functional. Testicular tissue is abnormal and sterile, but produces high levels of testosterone. This phenomenon also characterizes other talpid species from Europe and North America. To study the origin of this singular reproductive specialization, we examined the gonads of several female specimens belonging to two critical taxa. Although large Japanese moles (Mogera wogura) posses ovotestes, greater Japanese shrew moles (Urotrichus talpoides) are characterized by normal ovaries. The results fit parsimoniously with a recent phylogenetic study that places Urotrichus relatively basal in the talpid tree and separate from the American shrew mole. Parsimony reconstruction on alternative phylogenetic hypotheses clearly indicates that reversal(s) must have occurred and suggests that a relatively simple genetic mechanism must be associated with the evolution of female hermaphroditism in moles.

Early skeletal development in Talpa europaea, the common European mole.

An ontogenetic series of 22 cleared and double-stained prenatal specimens was used to study the sequence of ossification of selected postcranial skeletal elements of Talpa europaea. Results were compared with nine other therian mammals, with Alligator, Chelydra, and Lacerta as outgroups. Using the event-pairing method, shifts in the onset of ossification in T. europaea, Sus, and Homo were identified. In T. europaea, the ossification of the cervical vertebrae starts before the metatarsals. In Homo and Sus, the tarsals ossify before the pubic bone. These shifts in the sequence of ossification are unique among the mammals examined, whereas many other changes, characterising monophyletic groups and/or evolving convergently, were also identified. Particular attention was given to some peculiar calcified elements of the hand in T. europaea, which were identified as accessory ;sesamoid bones', and do not display a chondrified precursor. They start to calcify before all others of the hand and later fuse. They appear in all fingers and function as reinforcement for the distal phalanges, most likely as an adaptation for burrowing. The development of the sesamoid bones was examined using histological sections and macerated adults.

Early development of a somatosensory fovea: a head start in the cortical space race?

Star-nosed moles have 11 mechanosensory appendages surrounding each nostril, and primary afferents from a single appendage-the tactile fovea-are greatly over-represented in somatosensory cortex. It was found that the foveal appendage led development in the periphery, had the greatest innervated surface area in embryos, and developed mature nerve terminals and epidermal sensory organs first; also, in developing cortex, markers for metabolic activity (cytochrome oxidase) appeared first in the fovea representation. This developmental sequence may provide the fovea with an advantage in a competition for cortical space, and account for the much larger areas of cortex devoted to foveal afferents.

The development of a biological novelty: a different way to make appendages as revealed in the snout of the star-nosed mole Condylura cristata.

The nose of the star-nosed mole Condylura cristata is a complex biological novelty consisting of 22 epidermal appendages. How did this new set of facial appendages arise? Recent studies find remarkable conservation of the genes expressed during appendage formation across phyla, suggesting that the basic mechanisms for appendage development are ancient. In the nose of these moles, however, we find a unique pattern of appendage morphogenesis, showing that evolution is capable of constructing appendages in different ways. During development, the nasal appendages of the mole begin as a series of waves in the epidermis. A second deep layer of epidermis then grows under these superficial epidermal waves to produce 22 separate, elongated epidermal cylinders embedded in the side of the mole's face. The caudal end of each cylinder later erupts from the face and rotates forward to project rostrally, remaining attached only at the tip of the snout. As a result of this unique 'unfolding' formation, the rostral end of each adult appendage is derived from caudal embryonic facial tissue, while the caudal end of each appendage is derived from rostral facial tissue. This developmental process has essentially no outgrowth phase and results in the reversal of the original embryonic orientation of each appendage. This differs from the development of other known appendages, which originate either as outgrowths of the body wall or from subdivisions of outgrowths (e.g. tetrapod digits). Adults of a different mole species (Scapanus townsendii) exhibit a star-like pattern that resembles an embryonic stage of the star-nosed mole, suggesting that the development of the star recapitulates stages of its evolution.