Hand clasping--an overview]

The literature concerning the asymmetry clasping hands is reviewed based on 192 studies. This paper describes the incidence, sex differences, age differences and genetical problems including twinning. The incidence of left hand clasping ranges from 30% to 75% (mean 43%). The review confirms the so-called east-west-gradient and there is a predominance of the left type in Europe. Age and sex differences are only small. There is only a small relationship between hand clasping and handedness. 18 authors examined hand clasping in families and 4 in twins. The family data suggest that hand clasping may be under genetic control, yet it is clear that no simple genetic model for the inheritance can be applied. Both monozygotic and dizygotic twins show a low concordance and the R-R, R-L and L-L pairs in monozygotic and dizygotic twins are in binomial distribution.     

屯堡人7项不对称行为特征

在贵州省安顺市对359例屯堡人(男145例,女214例)中学生的7项不对称行为特征(扣手、利手、叠臂、叠腿、利足、起步类型和优势眼)进行了调查。研究结果显示:1)屯堡人除叠臂左型率高于右型率外,其他6项不对称行为特征均为右型率高于左型率;2)屯堡人7项不对称行为特征的出现率均无性别差异;3)与其他族群比较,屯堡人利足右型率和叠臂右型率中等,叠腿右型率和扣手右型率高;4)屯堡人两两不对称行为中,扣手与利手、扣手与叠臂、扣手与叠腿、叠腿与起步相关,利手、利足、起步三种行为彼此相关。     

Study on pottical type, palmar and plantar digital formulae, hand clasping, arm folding, handedness, leg folding and stride type in the Daur population, China

The pottical type, palmar and plantar digital formulae, hand clasping, arm folding, handedness, leg folding and stride type have been investigated on a sample of 143 male and 160 female students of the Daur population of Molidawa Banner, Inner Mongolia. The results of this study are the following: 1. the frequency of the hyperextensive pottical type is 49.17%, the relative length of index over annularis 12.21%, right hand clasping 45.87%, right arm folding 49.50%, right handedness 94.39%, right leg folding 72.28% and right stride type 44.88%, 2. pottical type, hand clasping, handedness, leg folding and stride type do not show significant sex differences, 3. there are some relations between hand clasping and arm folding as well as between arm folding and stride type, 4. compared with other population groups, the Daur population shows a low frequency of right hand clasping, a moderate frequency of right arm folding and a low frequency of left handedness.     

Hand clasping and arm folding in the Basque population

Hand clasping and arm folding have been analysed in 286 males and 455 females of the Basques. The results are compared with those from other Spanish populations. Examination of the association between hand clasping and arm folding shows a strong correlation between the right type of one and the left type of the other.     

中国木雅人、尔苏人、八甲人与空格人6项不对称行为的特征

对四川、云南尔苏人、木雅人、空格人、八甲人4个族群进行了6项不对称行为特征(扣手、利手、叠臂、利足、叠腿、起步)的调查,结果显示:1)4个族群中,仅尔苏人、木雅人、八甲人叠臂以L型出现率偏高以外,其余不对称行为特征均为R型出现率较高;2)4个族群6项不对称行为特征均不存在性别间差异,但木雅人、尔苏人与八甲人在扣手这一不对称行为特征上存在族群间差异,木雅人与空格人在叠臂这一不对称行为特征上存在族群间差异;3)统计分析了6项不对称行为特征之间的相关性,仅八甲人多数不对称行为特征之间存在相关性。     

Genetic associations between lateral signs.

Only a few authors examined several lateral asymmetries in families simultaneously. Seven lateralities (handedness, footedness, eyedness, earedness, hand clasping, arm folding and leg crossing) were examined in a sample of 292 biologically related parent-offspring triads and 36 sibling pairs. Analysis of the family data showed a significant association between parents and children in most lateralities. The frequency of left-sidedness increased with the number of left-sided parents. Only in the case of footedness and hand clasping there are no significant correlations. Additionally we investigated the inter-relationship between the lateralities in the parental generation. There is a relative strong relation especially among the functional asymmetries. The study analysed also the genetic association (linkage) according to McManus & Mascie-Taylor (1979) between seven different lateralized characteristics. It is shown that the lateral asymmetries correlate genetically relatively imperfectly with each other comparing the inter-relationship between the lateralities in the parental generation. This implicates a multigenetical and also multifactorial determination of laterality in human being.     

鄂伦春、鄂温克、达斡尔族一侧优势功能特征研究

报道了鄂伦春、鄂温克、达斡尔族7项一侧优势功能特征(扣手、利手、叠臂、叠腿、起步类型、优势足、优势眼)。结果显示,3个民族7项特征的左右型分布规律有较多的相似之处,绝大多数特征与性别无关。鄂温克族和达斡尔族的利手与叠腿、优势足、起步类型间均具明显关联,其腿足特征间的相关程度亦优于手臂特征。鄂伦春族则不然。 Abstract:Authors reported 7 traits of lateral functional dominance including hand clasping,handedness,arm folding,leg folding,stride type,foot preference and eye preference in Oroqen,Ewenki and Daur nationalities.The results showed as follws;(1)Among three nationalities their left and right distributive patterns of the traits had many similarities and there was no relation between most traits and sexes.(2)There were obvious corelations between handedness and leg folding,handedness and foot preference,handedness and stride type in Eweike and Daur nationalities.Compared with the traits of leg-foot with interrelation,the traits of hand-arm had little degree on correlation in Ewenki and Daur.(3)There was correlation between eye preference and hand clasping but no relation between handedness and hand clasping.     

A neurobehavioral model for rapid actions of corticosterone on sensorimotor integration.

Stress-induced corticosterone (CORT) secretion that causes a rapid blockade of courtship clasping by male roughskin newts (Taricha granulosa) is mediated by a specific neuronal membrane receptor for CORT. Amplectic clasping, which can be triggered by pressure on the ventral body surface and cloaca, is controlled by the influence of medullary neurons on the spinal cord. Using clasping as a simple neurobehavioral model, we have focused our analysis of CORT effects on clasping by examining the steroid's effects on neurophysiological properties of medullary neurons, especially medullary reticulospinal neurons, the principal output cells from the brain to the spinal cord. Systemic CORT caused, within 3 min of injection, diverse reductions in reticulospinal neuron excitability. Another rapid CORT effect on medullary neurons was to depress responsiveness to pressure on the cloaca. Experiments with chronically implanted, freely moving newts revealed that the rapid CORT effects are quite specific to neural processes related to clasping. CORT injections rapidly blocked clasping in response to cloacal stimuli and concurrently depressed neuronal responses to cloacal pressure and firing associated with clasping. Activity of reticulospinal neurons was often associated with nonclasping movements and this activity was rarely altered by CORT. Thus, CORT mainly affected aspects of neuronal function related to clasping. In other neurophysiological experiments, we found that the neuropeptides vasotocin and corticotropin-releasing hormone modified the neural effects of CORT. Prior exposure of medullary neurons to either of these neuropeptides caused systemic CORT administration to rapidly potentiate neuronal responses to cloacal stimuli, indicating that the direction and potency of CORT effects depend critically on the prevailing neuroendocrine state of the brain.     

Study on some genetic traits of Tibetans at Chandragiri, District Ganjam, Orissa

The incidence of ABH secretion, dry cerumen, ear-lobe attachment, hand clasping, arm-folding, and digital hair, occipital hair whorl and relative length of first and second toes is reported for Tibetan refugees. In those traits for which comparative data are available, they resemble Mongoloid rather than Caucasoid populations.     

The effects of sex steroids and vasotocin on behavioral responses to visual and olfactory sexual stimuli in ovariectomized female roughskin newts

Previous studies have found that vasotocin (AVT) administration to male roughskin newts (Taricha granulosa) enhances courtship clasping as well as appetitive responses to specific sexual stimuli and that treating female newts with androgens plus AVT induces the expression of male-typical courtship clasping (the selective clasping of females). However, the unique and/or interactive effects of sex steroids and AVT on appetitive responses to specific sexual stimuli have not yet been determined. To first identify male-typical, sexually dimorphic appetitive responses to female sexual stimuli, we tested intact newts during the breeding season and found that males, but not females, are attracted to female visual and pheromonal sexual stimuli. We then used ovariectomized (ovx) females implanted with empty silastic capsules (Blk) or with capsules containing testosterone (T), dihydrotestosterone (DHT), or estradiol (E2) and then injected with either saline or AVT to determine the effects of steroids and AVT, alone or in combination with each other, on male-typical behavioral responses to those stimuli. E2 treatment depressed responses toward female visual stimuli independently of AVT. On the other hand, only T-implanted, AVT-injected females displayed male-typical behavioral responses toward female olfactory stimuli, preferring to spend more time in proximity to female-scented than unscented newt models and selectively clasping the female-scented models. Together, these results support the conclusion that sex steroids and AVT influence behavioral responses to sexual stimuli via sensory-specific mechanisms. Furthermore, they suggest that T and AVT interact within the brain to influence sensorimotor processing in the pathways that integrate olfactory sexual stimuli into male-typical courtship behaviors.     

4 anthroposcopic markers in the Northern Greece population: hand folding, arm folding, tongue rolling and tongue folding

Four anthroposcopic traits, namely hand clasping, arm folding, tongue rolling and tongue curling have been studied in a total of 7763 individuals from Thessaloniki and its surroundings, representing a sample of the population of Northern Greece. The statistical analysis of the data indicates significant sex differences only concerning tongue rolling. The frequencies obtained for the four traits under study are compared with data from the literature.     

Hormone effects on male sex behavior in adult South African clawed frogs, Xenopus laevis

Intact, adult male Xenopus laevis were injected with human chorionic gonadotropin (HCG) and tested with intact HCG-primed females. Under these conditions, males displayed high levels of sex behavior (clasping of females). By 2 weeks after castration, these males had ceased clasping. Testosterone and dihydrotestosterone reinstated clasping in male castrates. Following removal of testosterone or dihydrotestosterone pellets, castrated males ceased to clasp. No male was ever observed to clasp following estradiol implanted in pellets or in silastic capsules. In experiments on castrated, adult, female Xenopus laevis, both testosterone and testosterone propionate pellets reliably produced male sex behavior in the form of clasping. The clasping of testosterone-implanted female and male castrates was very similar in form and duration. The behavioral effectiveness of testosterone in both sexes and the ineffectiveness of estradiol in eliciting clasping is paralleled by autoradiographic localization of sex steroids in brain where the distribution of testosterone-concentrating, cells is the same for males and females, but different from the distribution of estradiol-concentrating cells.     

汉族湘语族群的7项不对称行为特征

在湖南省娄底地区调查了汉族湘语族群507例(男269例、女238例)高中学生的7项不对称行为特征(扣手、利手、叠臂、叠腿、利足、起步类型和优势眼)。研究结果显示:1)湘语族群叠臂与起步类型L型率高于R型率, 其余5项均R型率高于L型率。2)湘语族群利手、叠臂、利足出现率存在性别间差异, 其余4项出现率均不存在性别间差异。3)在已发表的资料中, 湘语族群扣手和起步类型右型率中等偏高, 利手右型率处于中等水平, 叠臂和叠腿右型率低,利足和优势眼右型率高。4)主成分分析结果显示, 汉族各族群不对称行为特征有一定的共性, 相对接近于中国北亚类型族群, 而与南亚类型族群差距较大。中国族群间不对称行为特征的差异, 主要反映在叠臂率差异、利足率差异。     

Sexually dimorphic muscles in the forelimb of the Japanese toad,Bufo japonicus

During the breeding season, male anurans display clasping behavior by holding females with their forelimbs. This behavior is peculiar to males, and may require specializations in forelimb musculature. The present study revealed that five kinds of forelimb muscles were heavier in the male Japanese toad than in the female: the flexor carpi radialis (FCR), the flexor antibrachii medialis caput superius (FAMsup), the abductor indicis longus (AIL), the extensor carpi radialis caput superius (ECRsup), and the flexor antibrachii lateralis superficialis caput superius (FALSsup). In addition, one breast muscle, the coracoradialis (CR), was also heavier in males than in females. A quantitative analysis of muscle fibers processed for myosin ATPase activity showed that, in such “sexually dimorphic muscles” of the female, both fast (twitch) and slow (tonic) muscle fibers were of smaller diameter than in other forelimb muscles of both sexes (all male muscles plus “nondimorphic muscles” of the female). Moreover, both types of fibers were less numerous than in the corresponding muscles of the male. These results suggest that the “sexually dimorphic muscles” are used especially for clasping by the male and are degenerative or subnormal in the female. Slow muscle fibers were neither peculiar to, nor abundant in, these clasping muscles, although they may well be necessary for tonic and prolonged contractions of the forelimb muscles during clasping. The mechanism of sexual dimorphism may be a direct action of androgens on clasping muscles or an indirect action on clasping muscles via the innervating motoneurons.     

Male-Male Clasping May Be Part of an Alternative Reproductive Tactic in Xenopus laevis

Male Xenopus laevis frogs have been observed to clasp other males in a sustained, amplectant position, the purpose of which is unknown. We examined three possible hypotheses for this counter-intuitive behavior: 1) clasping males fail to discriminate the sex of the frogs they clasp; 2) male-male clasping is an aggressive or dominant behavior; or 3) that males clasp other males to gain proximity to breeding events and possibly engage in sperm competition. Our data, gathered through a series of behavioral experiments in the laboratory, refute the first two hypotheses. We found that males did not clasp indiscriminately, but showed a sex preference, with most males preferentially clasping a female, but a proportion preferentially clasping another male. Males that clasped another male when there was no female present were less likely to “win” reproductive access in a male-male-female triad, indicating that they did not establish dominance through clasping. However, those males did gain proximity to oviposition by continued male-male clasping in the presence of the female. Thus, our findings are consistent with, but cannot confirm, the third hypothesis of male-male clasping as an alternative reproductive tactic.     

Neuroendocrinology of context-dependent stress responses: vasotocin alters the effect of corticosterone on amphibian behaviors

The ability of an animal to respond with appropriate defensive behaviors when confronted with an immediate threat can affect its survival and reproductive success. In the roughskin newt (Taricha granulosa), exogenous corticosterone (CORT) rapidly blocks and vasotocin (VT) enhances reproductive behaviors (mainly clasping behavior). Electrophysiological studies have shown that pretreatment of male Taricha with VT counteracts the inhibitory effects of CORT on neuronal activity in the medulla. To test whether similar interactions between VT and CORT influence reproductive behaviors in Taricha, we recorded the time spent and incidence of clasping in males injected with VT or vehicle at 60 min and then CORT or vehicle at 5 min before presentation of a female. This study found that clasping behavior is suppressed in males that received vehicle and then CORT, but is not suppressed in males that received VT and then CORT. Considering these results and the possibility that the performance of clasping behaviors might cause increases in endogenous VT activity, we tested whether the suppressive effects of CORT administration on clasping behavior would occur in males that had recently clasped females. The study found that, in contrast to males that had been isolated from females, CORT administration did not suppress clasping behavior in males that had been allowed to clasp females for 60 min prior to the hormone injection. Our results suggest that, at least in this amphibian and perhaps in other animals, the neuroendocrine regulation of alternative behavioral responses to threats involves functional interactions between corticosteroids and VT-like peptides.     

Clasping behavior and the asymmetrically latitudinal structure of the testes in the male dwarf gourami Colisa lalia

During the course of spawning, the male dwarf gourami Colisa lalia exhibits a clasping behavior where the male bends his body around the female under the floating bubble nest. In this study, in order to investigate the role of this clasping behavior, we observed the mating behavior of C. lalia in aquaria and conducted examinations of the male’s reproductive system anatomically and histologically. Forcing two males to compete for spawning with a female resulted in one male becoming dominant over the other and ultimately clasping the female. The losing male did not attempt to “sneak” during spawning, and it was possible to confirm that the clasping behavior is indispensable for sperm emission and only performed by the dominant male. Although testes of teleosts generally longitudinally elongate (i.e., parallel to the body axis), the testes of C. lalia were latitudinally elongate (i.e., vertically with respect to the body axis) and located asymmetrically between the liver and gallbladder. The latitudinal structure of the testes was also confirmed histologically, as the main testicular duct develops vertically. The asymmetric testes did not appear to influence the way in which the body bent during the clasping maneuver because the male was observed to bend its body equally to the right and left during clasping. However, the testes were situated in the vicinity of the fulcrum formed when the male bends its body, suggesting that the latitudinal testes are structurally related to its clasping behavior (i.e., a mechanism analogous to that of a nutcracker). Following physical principles (i.e., levers and mechanical advantage), it can be proposed that male dwarf gourami squeeze semen out from their latitudinal testes by bending their bodies.     

Rapid, Corticosterone-Induced Disruption of Medullary Sensorimotor Integration Related to Suppression of Amplectic Clasping in Behaving Roughskin Newts (Taricha granulosa)

Endogenously secreted or injected corticosterone (CORT) rapidly suppresses courtship clasping in male roughskin newts (Taricha granulosa) by an action on a specific neuronal membrane receptor. Previous studies, using immobilized newts, showed that CORT administration rapidly depresses excitability of reticulospinal neurons and attenuates medullary neuronal responsiveness to clasp-triggering sensory stimuli. The present study used freely moving newts to examine clasping responses and concurrently record sensorimotor properties of 67 antidromically identified reticulospinal and other medullary reticular neurons before and after CORT injection. Before CORT, reticulospinal neurons fired in close association with onset and offset of clasps elicited by cloacal pressure. Reticulospinal neurons also showed firing correlates of nonclasping motor events, especially locomotion. Neuronal activity was typically reduced during clasping and elevated during locomotion. Medullary neurons that were not antidromically invaded (unidentified neurons) usually showed sensorimotor properties that resembled those of reticulospinal neurons. Intraperitoneal CORT (but not vehicle) reduced the probability and quality of hindlimb clasping in response to cloacal pressure, especially within 5–25 min of injection. Simultaneously, responses of reticulospinal and unidentified neurons to cloacal pressure and occurrence of clasping-related activity were attenuated or eliminated. CORT effects were relatively selective, altering clasping-related neuronal activity more strongly than activity associated with nonclasping motor events. The properties of CORT effects indicate that the hormone impairs clasping by depressing processing of clasp-triggering afferent activity and by disrupting the medullary control of clasping normally mediated by reticulospinal neurons. The rapid onset of these CORT effects implicates a neuronal membrane receptor rather than genomic action of the steroid.     

Sex steroids and vasotocin interact in a female amphibian (Taricha granulosa) to elicit female-like egg-laying behavior or male-like courtship.

Female egg-laying behaviors and male amplectic clasping behaviors in the rough-skinned newt (Taricha granulosa) are similar in that animals clasp an object. In the case of egg-laying, females clasp submerged inanimate objects, whereas in amplexus, males clasp conspecific females. Considering these behavioral similarities and differences, we investigated the possibility that gonadal steroids and vasotocin (AVT) interact to control egg-laying behaviors, as has been shown for the control of amplexus in Taricha males. Intact, gravid T. granulosa females injected ip with AVT, unlike those injected with saline, exhibited egg-laying behaviors and oviposition. In ovariectomy-steroid-implant studies, no saline-injected female exhibited egg-laying behaviors, whereas AVT-injected ovariectomized females exhibited egg-laying behaviors if implanted with estradiol (E2), testosterone, or dihydrotestosterone (DHT), and not if implanted with empty capsules. When given a choice between clasping aquatic vegetation or other females (amplectic clasping), following an AVT injection, unoperated and sham-operated control females and ovariectomized females with E2 implants did not preferentially clasp aquatic vegetation over other females. In contrast, AVT-injected ovariectomized females with DHT implants preferentially clasped other females. Thus, exposure of Taricha females to estrogens or androgens appears to determine whether the AVT-induced clasping is egg-laying or amplectic clasping.     

Genetic studies of Pangwalas, Transhumant and Settled Gaddis. 4. Colour blindness, mid-phalangeal hair, ear lobe attachment and behavioural traits

Frequency distributions of colour blindness, midphalangeal hair, ear lobe attachment, hand clasping, arm folding, leg folding and handedness are reported for different population groups from Himachal Pradesh, North India, namely Pangwalas, Transhumant Gaddis (Brahmans, Rajputs and Scheduled Castes) and Settled Gaddis (Brahmans, Rajputs and Schedules Castes). An attempt has been made to compare the results of the present study within and between these groups as well as with the results of other reports from different population groups of India and Asia.