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Human anatomical science and illustration: the origin of two inseparable disciplines.

From the early wall paintings of ancient Egyptians to the recent advent of computer graphics, medical illustrators have employed a variety of techniques and materials to enrich the art of medicine. Over the centuries, medical illustrators have captured the variety of physical findings observed in the clinical, surgical, or postmortem settings and transferred them to a permanent medium. Specifically, the study of human anatomy has enjoyed a historically popular courtship with medical artistry since 1543, when Andreas Vesalius published his now legendary work entitled De Humani Corporis Fabrica. However, the development and subsequent advancement of human anatomical illustration are indebted to individuals whose lifetimes span several centuries prior to Vesalius. The scientific achievements in anatomy manifest not only an advancement of knowledge, but also are a reflection of cultural, political, and religious beliefs. With respect to the development of human anatomic illustration, three elements were essential: the recognition of anatomy as a distinct branch of medical science, the acceptance of human dissection as a scientific method to advance understanding of anatomical structure, and the advancement in printing such that illustrations could be included alongside descriptive text. This brief study will examine these milestones while highlighting the origin of anatomical illustration in its historical context and its relationship to the development of human anatomy as a recognized medical science.

Anatomy↗

Mutation, selection and the heritability of complex traits.

It has been suggested that complex traits, like intelligence, have low heritabilities. This hypothesis stems from the idea that strong selection for higher intelligence has led to the fixation of genetic variation for this trait. The same hypothesis has been framed for complex sexual ornaments used in courtship display. These traits are also subject to directional selection, in this case caused by sexual selection. However, contrary to the hypothesis, comparative data shows that sexual ornaments have higher additive genetic variation than similar non-sexual traits. It appears that the number of variable genes and the effect per genetic locus have increased for sexual ornaments. Theory suggests this is due to selection for extreme phenotypes resulting in condition-dependent expression of sexual traits. Experimental work on the stalk-eyed fly, Cyrtodiopsis dalmanni, confirms that the male sexual trait (exaggerated eyespan) is more sensitive to environmental conditions than other non-sexual traits (wing dimensions and female eyespan). This environmental sensitivity has a genetic basis and environmental stress enhances genetic differences. It is likely that genetic variation in intelligence is maintained in a similar fashion.

Animals↗

Assortative marriage for physical characteristics in newlyweds.

The objective of this study was to estimate the degree of spouse resemblance in the first 6 months of marriage and to compare these estimates with those of previous studies on long-standing marriages. The participants, 215 couples, were selected from the marriage licenses issued in two counties within the San Francisco Bay Area. They attended one of several group sessions in which questionnaires were filled out and forearm length measurements were taken. Only the results for the couples of European ancestry (N = 164) are presented here since the other subsamples were too small to analyze separately. The husband-wife correlations for body size were found to be essentially equal to those in long-standing marriages. Thus, it appears that physical resemblance in couples is the result of initial assortment rather than convergence arising from long cohabitation. A significant residual husband-wife correlation for weight was found after statistically partialing out height. It may be that assortative marriage for food habits and attractiveness produces spouse resemblance in weight, independently of height. Alternatively, convergence in eating habits may occur during courtship and the early months of marriage which may increase the resemblance in weight. There was very little homogamy for eye color independently of race, and no evidence of assortative marriage for handedness.

Adolescent↗

The pheromonal role of cuticular hydrocarbons in Drosophila melanogaster.

Pheromones play a crucial role in mate stimulation and discrimination. In the fruit fly Drosophila, the most abundant cuticular hydrocarbons act as sex pheromones during courtship behavior. There are several active molecules and they compose a sex- and species-specific pheromonal bouquet. Different species from the Drosophila melanogaster subgroup have adopted alternative systems of chemical mate recognition. Recent exploration of these interspecific variations, and of intraspecific variations, has led to the characterization of genes and to the mapping of structures that process the production and perception of chemical messages.

Alkadienes↗

Vocal-acoustic circuitry and descending vocal pathways in teleost fish: convergence with terrestrial vertebrates reveals conserved traits.

Vocal behavior is multifaceted and requires that vocal-motor patterning be integrated at multiple brain levels with auditory, neuroendocrine, and other social behavior processes (e.g., courtship and aggression). We now provide anatomical evidence for an extensive vocal network in teleost fishes (Batrachoididae: Porichthys notatus; Opsanus beta) that is strongly integrated with neuroendocrine and auditory pathways and that exhibits striking similarities to the vocal-acoustic circuitry known for mammals. Biotin compound injections into neurophysiologically identified vocal regions of the forebrain (preoptic area and anterior hypothalamus) and of the midbrain (periaqueductal gray and paralemniscal tegmentum) reveal extensive connectivity within and between these regions, as well as reciprocal relationships with the auditory thalamus and/or auditory midbrain (torus semicircularis). Thus, specific components of the basal forebrain and midbrain are here designated as the forebrain vocal-acoustic complex (fVAC) and midbrain vocal-acoustic complex (mVAC), respectively. Biotin injections into the mVAC and a previously identified hindbrain vocal pattern generator likewise provide anatomical evidence for a distributed network of descending projections to the vocal pacemaker-motoneuron circuitry. Together, the present experiments establish a vocal-auditory-neuroendocrine network in teleost fish that links the forebrain and midbrain to the hindbrain vocal pattern generator (i.e., fVAC --> mVAC --> pattern generator) and provides an anatomical framework for the previously identified neuropeptide modulation of vocal activity elicited from the forebrain and midbrain, which contributes to the expression of sex- and male morph-specific behavior. We conclude with a broad comparison of these findings with those for other vertebrate taxa and suggest that the present findings provide novel insights into the structure of conserved behavioral regulatory circuits that have led to evolutionary convergence in vocal-acoustic systems.

Animals↗

Nitric oxide/cyclic guanosine monophosphate signaling in the central complex of the grasshopper brain inhibits singing behavior.

Grasshopper sound production, in the context of mate finding, courtship, and rivalry, is controlled by the central body complex in the protocerebrum. Stimulation of muscarinic acetylcholine receptors in the central complex has been demonstrated to stimulate specific singing in various grasshoppers including the species Chorthippus biguttulus. Sound production elicited by stimulation of muscarinic acetylcholine receptors in the central complex is inhibited by co-applications of various drugs activating the nitric oxide/cyclic guanosine monophosphate (cGMP) signaling pathway. The nitric oxide-donor sodium nitroprusside caused a reversible suppression of muscarine-stimulated sound production that could be blocked by 1H-[1,2,4]oxadiazolo-[4,3-a]quinoxaline-1-one (ODQ), which prevents the formation of cGMP by specifically inhibiting soluble guanylyl cyclase. Furthermore, injections of both the membrane-permeable cGMP analog 8-Br-cGMP and the specific inhibitor of the cGMP-degrading phosphodiesterase Zaprinast reversibly inhibited singing. To identify putative sources of nitric oxide, brains of Ch. biguttulus were subjected to both nitric oxide synthase immunocytochemistry and NADPH-diaphorase staining. Among other areas known to express nitric oxide synthase, both procedures consistently labeled peripheral layers in the upper division of the central body complex, suggesting that neurons supplying this neuropil contain nitric oxide synthase and may generate nitric oxide upon activation. Exposure of dissected brains to nitric oxide and 3-(5'hydroxymethyl-2'-furyl)-1-benzyl indazole (YC-1) induced cGMP-associated immunoreactivity in both the upper and lower division. Therefore, both the morphological and pharmacological data presented in this study strongly suggest a contribution of the nitric oxide/cGMP signaling pathway to the central control of grasshopper sound production.

Animals↗

Identification of roughskin newt medullary vasotocin target neurons with a fluorescent vasotocin conjugate.

Arginine8 vasotocin (AVT), a neurohypophyseal peptide in nonmammalian vertebrates, plays a key role in the regulation of social behaviors related to reproduction. In male roughskin newts (Taricha granulosa), AVT is an important facilitator of several reproductive behaviors, including courtship clasping of females. Although AVT is known to act in certain brain regions and AVT receptors have been localized to some extent, specific target neurons for this peptide have not been identified in any species. Internalization of a receptor-specific conjugate of AVT and the fluorescent dye Oregon green was used to identify AVT target cells in the medulla of male roughskin newts. Medullary neurons are of interest because they appear to mediate facilitation of clasping by AVT. Direct application of AVT-Oregon green to the fourth ventricular surface of the medulla in vivo resulted in conjugate internalization by a widespread population of medullary neurons, particularly in the medial reticular formation and nuclei of cranial nerves V, VII, VIII, IX, and X. Some fourth-ventricle ependymal cells were also labeled. Reticulospinal neurons, which play an important role in clasping, were identified by retrograde labeling with tetramethylrhodamine dextran amine. AVT-Oregon green was internalized by 72% of these neurons. These results show that AVT can directly affect a very large and diverse medullary neuronal population, which may underlie the peptide's role in multiple neuroendocrinological processes, including autonomic and behavioral regulation. Selectivity of the AVT action may arise through interactions between AVT and steroids such as corticosterone.

Animals↗

Hormone concentrating cells in vocal control and other areas of the brain of the zebra finch ( Poephila guttata).

Using the autoradiographic method in the zebra finch (poephila guttata), areas of the brain were identified which contain cells which accumulate testosterone. Among these areas are the caudal nucleus of the hyperstriatum ventrale, nucleus intercollicularis of the midbrain, and the tracheosyringeal portion of the hypoglossal nucleus of the medulla (nXIIts). These three are known to control or influence androgen dependent song and other vocalizations of passeriform birds, and nXIIts is composed of the motoneurons innervating the vocal (syringeal) muscles. Other areas containing hormone-concentrating cells are the medial preoptic area, nucleus periventricularis magnocellularis of the hypothalamus, dorsal infundibular layers, dorsomedial thalamus, lateral septum, magnocellular nucleus of the anterior neostriatum, periventricular medial neostriatum, nucleus taeniae of the archistriatum, and ventral paleostriatum augmentatum. Accumulation by cells in the preoptic area, hypothalamus, and limbic forebrain is consistent with a general vertebrate pattern of distribution of brain cells which accumulate sex steroids. Some of these same areas may be involved in the control of androgen dependent events such as courtship, copulation, aggression, and feedback regulation of the hypophysis.

Animals↗

Ontogeny and sexual dimorphism of the sonic motor nucleus in the oyster toadfish.

Male oyster toadfish, Opsanus tau, produce a courtship boatwhistle call and have a larger sound-producing organ than females, who do not boatwhistle . We investigated the possibility of sexual dimorphism in ontogeny of the sonic motor nucleus (SMN) of the toadfish. Brain weight increases for life though the increase decelerates with increasing fish weight. Neuron number, ranging from 760 to 2,888 in the SMN, increases rapidly to about 3 years, more slowly to about 8 years, and then levels off. There are no sexual differences in regressions of brain weight and SMN neuron number against fish size or age. Neuronal soma size in the SMN increases from 8 to 35 micron in average diameter and 67 to 916 micron2 in area over a period of at least 7 years. Males have larger neurons than females (P less than .01). However, males can be separated by inspection into populations with large and small soma sizes. Neuron size is not different between females and males with small somas. Neurons in males with large somas are larger than neurons in females and neurons in males with small somas. Such male dimorphism is reminiscent of other behavioral and morphological dimorphisms , which have led to characterization of males into territorial and satellite forms in certain mating systems.

Air Sacs↗

Sexual differentiation of androgen accumulation within the zebra finch brain through selective cell loss and addition.

In zebra finches males, but not females, respond to gonadal androgens with the production of a stereotyped courtship song. Corresponding to this dimorphism in hormone sensitivity, two song-related neural regions, magnocellular nucleus of the anterior neostriatum (MAN) and hyperstriatum ventralis pars caudalis (HVc), have more cells that accumulate androgens in males than in females. The early hormonal environment establishes these sex differences in the brain and behavior such that females treated with estradiol (E2) shortly after hatching have male-typical levels of androgen accumulation in MAN and HVc, and respond to adult androgen stimulation with male-typical song. Using autoradiographic procedures employing the androgen 3[H]-dihydrotestosterone (DHT), the present study focused on whether E2 masculinizes androgen target cell numbers in MAN and HVc by inducing, or rather preserving, a male-typical number of androgen target cells. First, it was demonstrated that E2 treatment of females as late as 20 days after hatching increases the number of MAN and HVc cells accumulating 3[H]-DHT or its metabolites in adulthood to levels typical of males. Then the number of androgen target cells present within these nuclei in 20-day-old females was compared with the number present in adult females previously masculinized with E2 treatment on day 20. In MAN, E2 masculinizes androgen accumulation by preserving androgen target cell numbers in the face of massive overall cell loss, a result that suggests that E2 promotes the survival of androgen target cells during sexual differentiation. In HVc, however, E2 treatment on day 20 promotes an addition of androgen target cells, suggesting that during adolescence, E2 regulates the proliferation, migration, and/or survival of these cells.

Animals↗

The motor neurons innervating the direct flight muscles of Drosophila melanogaster are morphologically specialized.

The anatomy of the motor neurons innervating six direct flight muscles in Drosophila melanogaster has been investigated by using a horseradish peroxidase backfilling technique. The somata of these motor neurons are arranged in two distinct clusters ipsilateral to the muscle they innervate. One cluster of cell bodies is located in the ventrolateral region between the prothoracic neuromere and the mesothoracic leg-related neuropil and the other is situated dorsally and posteriorly to the mesothoracic leg-related neuropil. Axons from somata in the ventrolateral cluster run in the anterior dorsal mesothoracic nerve, while axons from somata in the other cluster run in the mesothoracic accessory nerve. This distribution of somata and axons is discussed in the light of the morphological similarity and proximity of these functionally related muscles. On the basis of the branching patterns of their neurites, direct flight muscle motor neurons can be classified as stubbly, fibrous or tufted. The terminal arborizations of the motor neurons over the direct flight muscles are also morphologically specialized. Both the central and the peripheral morphological specializations of the direct flight muscle motor neurons correlate with the activity patterns exhibited by their associated muscles during flight and courtship song.

Animals↗

Sexual and seasonal differences in the vomeronasal epithelium of the red-backed salamander (Plethodon cinereus).

Sexually dimorphic behaviors often are associated with sexually dimorphic neural structures. Perinatal hormonal levels influence structural sexual dimorphism, and seasonal structural changes also can be the result of seasonal hormonal fluctuations. We compared the volume of vomeronasal organs of male and female red-backed salamanders (Plethodon cinereus) throughout the year. Odorants are delivered to vomeronasal receptors through nasolabial grooves when salamanders touch the bases of these grooves to objects (nose tapping). Males may locate and identify potential mates through nose tapping during the prolonged breeding season that lasts from October through May. We compared vomeronasal organ data through multiple regression by using total body size, sex, and season as variables that may influence vomeronasal organ volume. Gonads also were examined as an indicator of reproductive status. Total body size and sex significantly affect vomeronasal organ volume; as body size increases, so does vomeronasal organ volume, and males have significantly larger vomeronasal organs than females at all times of the year. During the summer, both males and females have larger vomeronasal organs than at any other time of the year. Summer also is a period of intense foraging and the initiation of a new cycle of gametogenesis. Previous studies of circulating hormone levels in amphibians have shown that the initial stages of gametogenesis correspond to a period of low estradiol and testosterone levels but high gonadotropin-releasing hormone levels. The functional significance of sexual and seasonal differences in the vomeronasal organs of P. cinereus may relate to the neurogenesis of specialized receptors for courtship and mating.

Animals↗

Response of sparrows to songs of deaf and isolation-reared males: further evidence for innate auditory templates.

Adult song sparrows (Melospiza melodia) were tested for response to songs of conspecific males that had been reared in acoustic isolation or deafened early in life. Territorial males responded more aggressively to playback of songs of isolated males than to songs of deafened males but did not discriminate between songs of deafened males and heterospecific songs. Captive females showed an identical pattern when their response was measured in terms of the number and intensity of courtship displays evoked by song. The results support the auditory template theory of song development in demonstrating that species-identifying features are present in songs of isolates and in showing that males unable to hear their own song production do not incorporate these species-identifying features into their songs.

Animals↗

Prenatal antagonism of stress by naltrexone administration: early and long-lasting effects on emotional behaviors in mice.

The effects of prenatal exposure to stress and to naltrexone on emotional behaviors were studied in CD1 mice during ontogeny and in the adulthood. During ontogeny (a) lower body weights were initially found in pups born by mothers injected with naltrexone; (b) treatments did not affect sensory motor development except in the case of the cliff aversion reflex which occurred earlier in pups prenatally exposed to stress; (c) measures of ultrasonic vocalizations in stressful context showed that the amount of vocalizations emitted by pups born by stressed mothers was significantly higher than that emitted by pups born by naltrexone injected and control mothers (d) an examination of mother-offspring interactions on the very first day of observation indicated a consistent trend in stressed mothers to be more responsive to their pups. In adulthood, ultrasonic calls in courtship after short and long periods of isolation showed a time-dependent decrease of vocalizations in males prenatally exposed to naltrexone. These results indicate that the modifications of emotionality evident during early development are directly related to the reactivity of the mothers to the experimental treatments.

Animals↗

Motor training and physical fitness: possible short- and long-term influences on the development of individual differences in behavior.

Individual differences in the behavior of young and adult animals have been documented in diverse species. Possible sources of such variation are of interest to scientists representing many disciplines, including behavior, genetics, and population and evolutionary biology. Two variables that may be important in the ontogeny and maintenance of behavioral differences are (1) individual physical (aerobic and anaerobic) fitness and (2) possible genetic variations underlying individual abilities to engage in, and to benefit from, motor training early in life. The differential development of aerobic and anaerobic capacities may play a significant role in the ontogeny of individual differences in the performance of various motor skills. There also may be short- and long-term consequences of variations in physical fitness that influence individual abilities to perform energy demanding acts during aggressive encounters, interactions with prey or predators, and courtship and breeding. Genetic studies of a limited number of species indicate that specific genotypes are correlated with individual variations in motor performance, even among conspecifics. Multidisciplinary research concerning possible relationships among the ontogeny of physical fitness, genetics, and variations in behavior is needed. Recent work on the relationship between individual differences in physical fitness and variations in the behavior of adult cold-blooded vertebrates provides a good model for comparative research on warm-blooded species.

Animals↗

Ontogeny of sexually dimorphic ultrasonic vocalizations in Mongolian gerbils.

Sexual differentiation of emission rates (Experiment 1) and physical structure of ultrasonic vocalizations (Experiment 2) were investigated in Mongolian gerbils between 17 and 85 days of age. Animals from different litters were allowed to interact in iso- and heterosexual pairs. Vocalization rates increased in all groups, reaching a plateau at approximately day 56. Female pups had significantly higher rates of all vocalizations than male pairs before the plateau stage. Five stereotypic categories of physical structure were easily distinguished and confirmed by sound spectrographic analysis. Only mid-range frequencies of vocalizations appeared to decline throughout the juveniles' lives: this change may communicate the animal's age. Two male and one female categories were sexually dimorphic and similar in structure to those made during courtship interactions. Rates of male-type vocalizations appear to increase as testicular androgens start rising.

Aging↗

Distribution of sex steroid hormone receptors in the avian brain: functional implications for neural sex differences and sexual behaviors.

Developmental and seasonal changes in the production of androgens, estrogens, and progestins seem to control sex-specific differentiation and seasonal changes in appetitive and consummatory sexual behaviors of birds. This results in profound sex differences in the quality (sex-specific) or quantity (sex-typical) of behaviors such as courtship, territoriality, or copulation. Steroids affect the brain by binding to intracellularly located receptors. The same brain areas express androgen, estrogen, and progesterone receptors in male and female brains. Sex differences in these genetically determined patterns occur in the size of neuron populations that intrinsically express sex steroid receptors. Further permanent sex differences are subsequent to degenerative fates of receptor expressing neuron populations during ontogeny. Transient sex differences in receptor expression appear to be due to area-specific up- and down-regulation of receptor levels, reflecting transient changes in the level of circulating steroids, changes in environmental conditions, or in the physiological status of the individuals. In particular, intrinsic sex differences in the expression pattern of sex steroid receptors and steroid-independent regulation of the expression level of these receptors in the brain are limiting mechanisms for gonad-dependent sexual development and activities.

Animals↗

Rectal gland of Bactrocera papayae: ultrastructure, anatomy, and sequestration of autofluorescent compounds upon methyl eugenol consumption by the male fruit fly.

Sexually mature males of Bactrocera papayae are strongly attracted to and consume methyl eugenol (ME). Upon consumption, ME is biotransformed to two phenylpropanoids, 2-allyl-4,5-dimethoxyphenol (DMP) and (E)-coniferyl alcohol (CF), that are transported in the hemolymph, sequestered and stored in the rectal glands, and subsequently released as sex and aggregation pheromones during courtship. To date, very little work on the ultrastructure and anatomy of the rectal gland has been done, and the accumulation of phenylpropanoids in the rectal glands of males has not been observed visually. Our objectives are to describe the anatomy and fine structures of the rectal glands of males and females and to observe the accumulation of autofluorescent compounds in the rectal glands of males. The rectal glands of males and females have four rectal papillae with each papilla attached to a rectal pad. The rectal pads protrude from the rectal gland as the only surfaces of the gland that are not surrounded by muscles. The rectal papillae of ME-fed males had oil droplets and autofluorescent compounds that were absent from those of ME-deprived males. The autofluorescent compounds accumulated in the rectal sac, which is an evagination that is not found in rectal glands of females. The accumulation of these compounds increased with time and reached maximum at a day post-ME feeding and decreased thereafter. This trend is similar to the accumulation pattern of phenylpropanoids, CF and DMP in the rectal gland.

Animals↗