Selection on bristle length has the ability to drive the evolution of male abdominal appendages in the sepsid fly Themira biloba.

Many exaggerated and novel traits are strongly influenced by sexual selection. Although sexual selection is a powerful evolutionary force, underlying genetic interactions can constrain evolutionary outcomes. The relative strength of selection vs. constraint has been a matter of debate for the evolution of male abdominal appendages in sepsid flies. These abdominal appendages are involved in courtship and mating, but their function has not been directly tested. We performed mate choice experiments to determine whether sexual selection acts on abdominal appendages in the sepsid Themira biloba. We tested whether appendage bristle length influenced successful insemination by surgically trimming the bristles. Females paired with males that had shortened bristles laid only unfertilized eggs, indicating that long bristles are necessary for successful insemination. We also tested whether the evolution of bristle length was constrained by phenotypic correlations with other traits. Analyses of phenotypic covariation indicated that bristle length was highly correlated with other abdominal appendage traits, but was not correlated with abdominal sternite size. Thus, abdominal appendages are not exaggerated traits like many sexual ornaments, but vary independently from body size. At the same time, strong correlations between bristle length and appendage length suggest that selection on bristle length is likely to result in a correlated increase in appendage length. Bristle length is under sexual selection in T. biloba and has the potential to evolve independently from abdomen size.

Distribution of cells projecting to thalamus vs. those projecting to cerebellum in subdivisions of the dorsal column nuclei in raccoons.

To learn the distribution of cells projecting to the thalamus, as opposed to the cerebellum, in the mechanosensory nuclei of the dorsal medulla of raccoons, we analyzed the retrograde transport of horseradish peroxidase from the ventrobasal complex of the thalamus and from the cerebellum. We found six nuclear regions projecting heavily to the thalamus with very small projections to the cerebellum: Bischoff's, central cuneate, central gracile, rostral cuneate, rostral gracile nuclei, and cell group z. Two regions showed heavy projections to the cerebellum with no projections to the thalamus: the lateral portion of the external cuneate nucleus and the compact portion of cell group x. Four regions showed more equivalent projections to both target regions: basal cuneate, medial portion of the external cuneate nucleus, medial tongue extension of the external cuneate nucleus, and reticular portion of cell group x. Three more ventral regions were labeled: lateral cervical nucleus from thalamic injections but not from cerebellar injections; central cervical nucleus from cerebellar injections, which crossed the midline, but not from thalamic injections; and lateral reticular nucleus from both target regions. In most medullary regions, most cells project to one target and very few project to the other; we suggest that the cells projecting to the minor target convey samples of the information going to the major target.

Kinesthetic cortical area anterior to primary somatic sensory cortex in the raccoon (Procyon lotor).

Extracellular microelectrode recording of cortical unit activity, with subsequent histological examination, was used to determine the extent, organization, and cytoarchitecture of the zone of muscle afferent projections (kinesthetic cortex) anterior to the primary somatic sensory cortex in anesthetized raccoons. Activity was evoked in response to mechanical stimulation of muscles from which the overlying skin had been dissected away. Most kinesthetic responses were elicited in a contiguous cortical area, which included: the anterior bank of the lateral arm, and the fundus and posterior bank of the medial arm of the medial central sulcus; and the anterior two-thirds of the interfundic rise within the interbrachial sulcus. Some responses were recorded in a separate small area of the anterior bank at the medial end of the lateral central sulcus. Somatotopy was evident with forelimb represented lateral to hindlimb. Proximal limb muscles were represented in the center of the medial central sulcus; distal muscle projections were medial (hindlimb) or lateral (forelimb) in the same sulcus. Most representations were of flexor and extensor muscles of the contralateral carpus and forepaw digits. Activity at a given recording locus in the kinesthetic area could be elicited by both flexor and extensor muscles, which acted about a common joint. Low amplitude units evoked by cutaneous stimulation of the dissected skin were recorded in the kinesthetic area; these were from receptive fields of skin that normally overlay the muscles whose higher-amplitude evoked kinesthetic units were represented in that same recording locus. The kinesthetic zone was anterior to primary somatic sensory cortex, where the outer stripe of Baillarger and granular layer IV become attenuated. In the hindlimb muscle representation area, the additional criterion of area 3a (large pyramidal cells in layer V) was seen. However, no cytoarchitecture could be identified that was consistently associated with the kinesthetic cortex.

Organization of intracortical and commissural connections in somatosensory cortical areas I and II in the raccoon.

The organization of intracortical and callosal projecting cell bodies was examined in somatosensory representation areas I (SI) and II (SII) of the raccoon by use of horseradish peroxidase (HRP) or horseradish peroxidase-wheat germ agglutin (HRP-WHA). HRP and HRP-WHA were injected into commissurally and noncommissurally connected subdivisions of SI and SII. Injection sites in SII were identified electrophysiologically. Results were obtained from transverse sections in which the HRP was visualized with the aid of the substrates dihydrochlorobenzidine or tetramethyl benzidine in the presence of hydrogen peroxidase. The principal findings were the following: (1) there are reciprocal connections between SI and SII; (2) in SI the intracortically projecting cell bodies and terminals are located primarily in sulcal cortex; (3) intracortically projecting neurons in SI are located primarily in layers III whereas in SII they are located principally in layers III and V; (4) there are connections between disparate areas within SI; and (5) there are intracortical connections between callosum-connected and acallosal regions in SII. These results are discussed with regard to the results of mapping studies of the SI, the significance of intracortical connections to the formation of sulci in SI, and the possible roles of nonhomotopic connections in the intermanual transfer of learning.

Medullary sources of projections to the kinesthetic thalamus in raccoons: external and basal cuneate nuclei and cell groups x and z.

In raccoons and other mammals, a pathway for kinesthetic sensation (from muscles, fascia, tendons, and joints) reaches the anterodorsal cap of the ventrobasal thalamus and the anteriormost part of the somatic sensory cerebral cortex. To find the medullary component of this kinesthetic pathway in raccoons, small injections of horseradish peroxidase were made in the thalamus under guidance of simultaneous electrophysiological recording from kinesthetic projections. As determined by retrograde labeling following these injections, kinesthetic thalamic subregions receive projections as follows: caudomedial from cells in the external cuneate nucleus and its medial tongue, rostromedial from cells in basal cuneate nucleus, and rostrolateral from cells in cell group z and the reticular division of cell group x. Electrophysiological recording showed kinesthetic representations in each of these medullary regions. Labeled cells were also observed in the infratrigeminal subnucleus of the lateral reticular nucleus. Cats have kinesthetic projections to the thalamus from the basal cuneate and cell group z; raccoons (and monkeys) have these plus projections from the external cuneate and cell group x. This suggests that the kinesthetic projection system in raccoons and monkeys is expanded in correlation with their more dextrous use of the hand.

Species differences in mechanosensory projections from the mouth to the ventrobasal thalamus.

To determine whether the largely ipsilateral, inverted representation of mouth parts in the ventrobasal thalamus of sheep was unique to that species or an expansion of a general mammalian pattern, the corresponding thalamic projections were mapped electrophysiologically in a selected series of mammals (oppossums, agoutis, squirrel monkeys, cats, raccoons, and sheep) representing major branches of evolution among therian mammals. In mapping, tungsten microelectrodes were used to record multi-unit discharges in the thalamus in response to mechanical stimulation of oral surfaces. The pattern of projections seen in sheep is not a general mammalian pattern; there is extensive variability among mammals in the laterality and internal orgainzation of the projections from the mouth. In spite of the great variability, the results suggest an hypothesis concerning phylogenetic trends: descendants of palaeoryctoid insectivores (cats, raccoons, and sheep in our sample) have extensive ipsilateral projections from the mouth, in other therian mammals (opossums, agoutis, and squirrel monkeys in our sample) the ipsilateral component is small or absent.

Organization of postcranial kinesthetic projections to the ventrobasal thalamus in raccoons.

To determine the presence and organization of kinesthetic, as compared with other mechanosensory projection zones in the thalamus of raccoons, unit-cluster responses to mechanical stimulation of the postcranial body were mapped electrophysiologically in the thalami of 14 raccoons anesthetized with Dial-urethane. A distinct zone of kinesthetic projections (from receptive fields in muscles, tendons, and joints) was found in the rostral and dorsal aspects of the mechanosensory projection zone. These projections are somatotopically organized: those from axial structures lie dorsalmost and those from successively more distal limb regions are successively more caudoventral. The kinesthetic forelimb representation is large and lies rostrodorsal to a large central core of cutaneous projections from the forepaw digits. A few scattered kinesthetic projections were found at the caudal edge of the sensory thalamic region. The large, spatially and somatotopically distinct kinesthetic projection zone in the thalamus parallels those seen in the cortex and medulla of raccoons. Similar findings in monkeys, and suggestions from data in cats and humans support the hypothesis of a distinct pathway to the cortex for kinesthetic information in all mammals.

Demarcations of the mechanosensory projection zones in the raccoon thalamus, shown by cytochrome oxidase, acetylcholinesterase, and Nissl stains.

To determine anatomically the boundaries and internal organization of the kinesthetic and cutaneous mechanosensory regions of the ventrobasal thalamus, alternate section series from electrophysiologically mapped tissues from 14 raccoons were stained for cytochrome oxidase, myelinated fibers, acetylcholinesterase, and Nissl substance. Microelectrode tracks, along with electrolytic lesions placed as tissue markers, reveal that the mechanoreceptor projection zones have higher cytochrome oxidase and lower acetylcholinesterase staining than some neighboring regions. Both these enzymatic stains reveal particularly sharp boundaries separating the mechanoresponsive region, from the lateral posterior nucleus dorsally and from the ventroposterior inferior nucleus ventrally. The kinesthetic projection zone is often separated from other mechanoreceptor projections by bundles as well as laminae of myelinated fibers, similar to those separating cutaneous projections from distinct body parts. These subdivisions are particularly well marked by the cytochrome oxidase stain. The combination, in neighboring sections, of the use of the several stains adds considerably to the visible delineation of these functionally distinct regions, beyond what can be seen in Nissl-stained sections.

The anterior border zones of primary somatic sensory (S1) neocortex and their relation to cerebral convolutions, shown by micromapping of peripheral projections to the region of the fourth forepaw digit representation in raccoons.

In raccoons the somatic sensory neocortex is greatly expanded, with separate gyral crowns devoted to and intervening sulci separating, sensory representations of separate body parts, most strikingly those of the volar surfaces of individual forepaw digits. Most of the cortex in this region is buried in widely ramifying sulcal walls, wherein sensory projections have not been studied. We have determined mechanosensory projections to the fourth digit representation region including all neighboring sulcal walls, using tungsten microelectrodes for 3-dimensional micromapping. We found no significant alteration in the location and pattern of projections when the following different anesthetics were used: dial-urethane, chloralose, or methoxyflurane with nitrous oxide. The precisely organized somatotopic representation of the distal volar surface of the fourth digit, on the causal aspect of its gyral crown, continues down the anterior bank of the triradiate sulcus. This meets, at the fundus, projections from the proximal volar surface of the digit which occupy the posterior sulcus wall; they in turn meet projections from the volar palm at the gyral crown. In the anterior part of the crown containing the representation of the distal volar digit, across the crown. In the anterior part of the crown containing the representation of the distal volar digit, across the crown of the gyral bridge intervening between the medial and lateral segments of the central sulcus, throughout the posterior walls of the central sulci, and in the walls of the interbrachial sulcus, we found a distinctive border-zone of projections from heterogeneous receptive fields. Within a roughly somatotopic basic pattern of organization we found intermingled projections from single and multiple claws and dorsal hairy surfaces of digits and proximal hand, along with additional projections from volar surfaces. These projections can be construed as forming something of a distorted mirror-image of the representation of the volar hand. Beyond this was a second zone of distinctive projections from afferents of the forelimb muscles, in the anterior walls of the central sulci. These projections are interrupted where the sulci are interrupted. The zone of muscle afferent projections corresponds to those seen between sensory and motor regions in other species; its strict association with sulcal folding here and in other species suggests a general relationship of these projections to central sulci. The zone of heterogeneous projections resembles similar zones seen at other levels of this system in raccoons, in the cortex of other species, and it may relate to some of the multiple representation reported in other species. It also may be related to the formation of sulci in this region and may be a specialized zone for cortico-cortical connections.

Mechanosensory projections to cuneate, gracile, and external cuneate nuclei in a tree squirrel (fox squirrel, Sciurus niger).

The distribution and organization of mechanosensory projections to the cuneate, gracile and external cuneate nuclei were mapped in tree squirrels anesthetized with ketamine and urethane. Tungsten microelectrodes were used to record unit and unit cluster responses to mechanical stimulation. Most responses in the gracile nucleus were to stimulation of skin and hairs of the tail, hind foot and leg, and trunk, in that order going from dorsal to ventral in the nucleus. A similar distribution of responses was found in the cuneate nucleus to stimulation of forelimb, neck and pinna going from dorsomedial to ventrolateral in the nucleus. Some responses to stimulation or movement of subcutaneous tissue were found in the cuneate and gracile nuclei. Responses in the external cuneate were to stimulation of deep-lying tissues in the hand medially, the lower arm centrally and the shoulder and trunk laterally. The pattern of projections in the tree squirrel differed most strikingly from that seen in raccoons and opossums in the relatively small extent of projections from the glabrous skin of the forepaws which were concentrated in a small region near the obex in squirrels. In contrast, there was a large representation of hairy receptive fields located on the forelimb in all regions of the squirrel cuneate nucleus. Otherwise, the somatotopic organization of mechanosensory projections to these dorsal column nuclei in tree squirrels was similar to that reported in other mammals.

Magnetic resonance imaging and three-dimensional reconstructions of the brain of a fetal common dolphin, Delphinus delphis.

To demonstrate the kinds of data that can be obtained non-destructively and non-invasively from preserved museum specimens using modern imaging technology the head region of a whole body fetal specimen of the common dolphin, Delphinus delphis, aged 8-9 months post-conception, was scanned using Magnetic Resonance Imaging (MRI). Series of scans were obtained in coronal, sagittal and horizontal planes. A digital three-dimensional reconstruction of the whole brain was prepared from the coronal series of scans. Sectional areas and three-dimensional volumes were obtained of the cerebral hemispheres and of the brainstem-plus-cerebellum. Neuroanatomical features identified in the scans include the major sulci of the cerebral hemispheres, well-differentiated regions of gray and white matter, the mesencephalic, pontine, and cervical flexures, the "foreshortened" appearance of the forebrain, and the large auditory inferior colliculi. These findings show that numerous features of the fetal common dolphin brain can be visualized and analyzed from MRI scans.

Influence of intonation on auditory sequential memory skills.

This study was designed to investigate the effect of intonation on the auditory sequential memory spans of children normal in language development and those with delay. Analysis indicated that intonation as a cue did not facilitate recall of monosyllabic nouns presented in sequences of two, three, four, and five words. Children with a language delay of six months or longer had spans of one word less than children with no language delay. Neither recall nor the memory span was related to chronological age over the range of 42 and 72 mo. for either group of children.