Hand biomechanics during simulated stone tool use

Human radial digits have derived features compared with apes, with long robust thumbs, relatively larger joint surfaces, and hypertrophic thenar muscles. Here we test the hypothesis that these features evolved in the context of making and using stone tools, specifically for producing large gripping forces and for countering large joint contact stresses. We used portable force plates simulating early stone tools to: 1) document and compare the magnitude of external/internal forces and joint stresses in the radial digits during hardhammer percussion and flake use, and 2) examine how variation in digit morphology affects muscle and joint mechanics during stone tool use. Force and kinematic data were collected from a sample representing normal variation in digit morphology (n = 25). The effects of digit size/shape on digit biomechanics were evaluated using partial correlations, controlling for tool reaction forces and impact velocities. Results show that individuals with longer digits require relatively less muscle force to stabilize digital joints, and are exposed to relatively lower joint contact stresses during stone tool use, due in part to an increase in the robusticity of metacarpals and phalanges in humans relative to chimpanzees. These analyses further suggest that Pan- or australopith-like pollical anatomy presents serious performance challenges to habitual tool use. Our data support the hypothesis that evolutionary increases in thumb length, robusticity, and thenar muscle mass enabled Homo to produce more force and to tolerate higher joint stresses during tool use.     

The developmental evolution of avian digit homology: An update

The identity of avian digits has been unresolved since the beginning of evolutionary morphology in the mid-19th century, i.e. as soon as questions of phylogenetic homology have been raised. The main source of concern is the persistent discrepancy between anatomical/paleontological and embryological evidence over the identity of avian digits. In this paper, recent evidence pertaining to the question of avian digit homology is reviewed and the various ideas of how to resolve the disagreement among developmental and phylogenetic evidence are evaluated. Paleontological evidence unequivocally supports the hypothesis that the fully formed digits of maniraptoran theropods are digits DI, DII, and DIII, because the phylogenetic position of Herrerasaurus is resolved, even when hand characters are excluded from the analysis. Regarding the developmental origin of the three digits of the avian hand the discovery of an anterior digit condensation in the limb bud of chickens and ostriches conclusively shows that these three digits are developing from condensations CII, CIII, and CIV. The existence of this additional anterior condensation has been confirmed in four different labs, using four different methods: Alcian blue staining, PNA affinity histochemistry, micro-capillary regression and Sox9 expression. Finally, recent evidence shows that the digit developing from condensation CII has a Hox gene expression pattern that is found in digit DI of mice forelimb and chick hind limbs. The sum of these data supports the idea that digit identity has shifted relative to the location of condensations, known as Frame Shift Hypothesis, such that condensation CII develops into digit DI and condensation CIII develops into digit DII, etc. A review of the literature on the digit identity of the Italian Three-toed Skink or Luscengola (Chalcides chalcides), shows that digit identity frame shifts may not be limited to the bird hand but may be characteristic of “adaptive” digit reduction in amniotes (sensu Steiner, H., Anders, G., 1946. Zur Frage der Entstehung von Rudimenten. Die Reduktion der Gliedmassen von Chalcides tridactylus Laur. Rev. Suisse Zool. 53, 537–546) in general. In this mode of evolution two digits are lost, in the course of the adaptation of the three anterior digits to a function that does not require the two posterior digits. This evidence suggests that the evolution of digits in tetrapods can proceed at least on two distinct levels of integration, the level of digit condensations and that of adult digits.     

Dehydration and cold hardiness in the Arctic Collembolan Onychiurus arcticus Tullberg 1876

Specimens of the Arctic Collembolon Onychiurus arcticus were exposed to desiccation at several subzero temperatures over ice and at 0.5 °C over NaCl solutions. The effects of desiccation on water content (WC), body fluid melting point (MP), supercooling point (SCP) and survival were studied at several acclimation temperatures and relative humidities. Exposure to temperatures down to −19.5 °C caused a substantial and increasing dehydration. At the lowest exposure temperature unfrozen individuals lost 91.6% of the WC at full hydration but more than 80% of the individuals survived when rehydrated. Exposure at 0.5 °C to decreasing relative humidities (RH) from 100% to 91.3% caused increasing dehydration and increasing mortality. Survival of equally dehydrated individuals was higher at subzero temperatures than at 0.5 °C. Concurrent with the decline in WC a lowering of the MP was observed. Animals exposed to −3 °C and −6 °C over ice for 31 days had a MP of −3.8 and < −7.5 °C, respectively. Specimens from a laboratory culture had a mean SCP of −6.1 °C, and acclimation at 0 or −3 °C had little effect on SCPs. Exposure at −8.2 °C over ice for 8 days, however, caused the mean SCP to decline to −21.8 °C due to the severe dehydration of these individuals. Dehydration at 0.5 °C in 95.1 and 93.3% RH also caused a decline in SCPs to about −18 °C. Individuals that had been acclimated over ice at −12.4 °C or at lower temperatures apparently did not freeze at all when cooled to −30 °C, probably because all freezeable water had been lost. These results show that O. arcticus will inevitably undergo dehydration when exposed to subzero temperatures in its natural frozen habitat. Consequently, the MP and SCP of the Collembola are substantially lowered and in this way freezing is avoided. The increased cold hardiness by dehydration is similar to the protective dehydration mechanism described in earthworm cocoons and Arctic enchytraeids. Accepted: 5 January 1998     

Role of RhoC in digit morphogenesis during limb development

Here we report a new role for the small GTPase RhoC in the control of limb chondrogenesis. Expression of rhoC is a precocious marker of the zeugopodial and digit blastemas and is induced by treatments with TGFbetas preceding the formation of ectopic digits. As development progresses, expression of rhoC outlines the growing distal tip of the digits, and marks the regions of interphalangeal joint formation. Functional experiments show that RhoC is a negative regulator of chondrogenesis, which controls digit outgrowth and joint segmentation. These functions appear to be mediated by reorganization of the actin cytoskeleton and modification of the adhesive properties of the mesenchymal cells.     

Evolution of digit identity in the three-toed Italian skink Chalcides chalcides: a new case of digit identity frame shift

SUMMARY Digit identity in the avian wing is a classical example of conflicting anatomical and embryological evidence regarding digit homology. Anatomical in conjunction with phylogenetic evidence supports the hypothesis that the three remaining digits in the bird wing are digits 1, 2, and 3. At the same time, various lines of embryological evidence support the notion that these digits develop in positions that normally produce digits 2, 3, and 4. In recent years, gene expression as well as experimental evidence was published that supports the hypothesis that this discrepancy arose from a digit identity shift in the evolution of the bird wing. A similar but less well-known controversy has been ongoing since the late 19th century regarding the identity of the digits of the three-toed Italian skink, Chalcides chalcides . Comparative anatomy identifies these digits as 1, 2, and 3, while embryological evidence suggests their derivation from embryological positions 2, 3, and 4. Here we re-examine this evidence and add gene expression data to determine the identity of the three digits of C. chalcides . The data confirm that the adult and the embryological evidence for digit identity are in conflict, and the expression of Hoxd11 suggests that digits 1, 2, and 3 develop in positions 2, 3, and 4. We conclude that in C. chalcides , and likely in its close relatives, a digit identity frame shift has occurred, similar to the one in avian evolution. This result suggests that changes in of digit identity might be a more frequent consequence of digit reduction than previously assumed.     

Claw retraction and protraction in the carnivora: Skeletal microvariation in the phalanges of the Felidae

All carnivorans retract and protract their claws. In felids and some viverrids the claws of digits II through V of both the manus and pes have a larger arc of rotation than those of other carnivorans; the claws retract to the lateral side of the middle phalanx rather than onto its dorsal surface as in most other carnivorans. This condition should be termed hyper-retraction. Morphological features of the middle and distal (ungual) phalanges that have been purported to be necessary for hyper-retraction in felids vary considerably among digits within the manus and pes. These features include the lateral projection of the distal head and the asymmetry of the shaft of the middle phalanx, and the oblique orientation of the articular surface on the distal phalanx. None of these features is necessary in every instance for hyperretraction, and some of the variation in these features is associated instead with protraction. Differences among digits in the orientation of the articular surface on the distal phalanx are associated with differences in the degree to which the claws must move laterally to rotate from the protracted to the retracted position. Differences in the orientation of the distal head on the middle phalanx are associated with the spreading of the claws during protraction. The manual claws are hook-shaped, whereas the pedal claws are more blade-like; this morphological difference is associated with differences in function between the manus and pes. In the manus the medial claws have a larger radius of curvature and a smaller angle of arc as compared to the more lateral claws; in the pes, the claws on digits III and IV have larger radii of curvature and smaller angles of arc. Digit I of the manus lacks the hyper-retraction mechanism; nonetheless, this digit shares many of the attributes that are associated with this mechanism. © 1996 Wiley-Liss, Inc.     

One Digit Interruption: The Altered Force Patterns during Functionally Cylindrical Grasping Tasks in Patients with Trigger Digits

Most trigger digit (TD) patients complain that they have problems using their hand in daily or occupational tasks due to single or multiple digits being affected. Unfortunately, clinicians do not know much about how this disease affects the subtle force coordination among digits during manipulation. Thus, this study examined the differences in force patterns during cylindrical grasp between TD and healthy subjects. Forty-two TD patients with single digit involvement were included and sorted into four groups based on the involved digits, including thumb, index, middle and ring fingers. Twelve healthy subjects volunteered as healthy controls. Two testing tasks, holding and drinking, were performed by natural grasping with minimal forces. The relations between the force of the thumb and each finger were examined by Pearson correlation coefficients. The force amount and contribution of each digit were compared between healthy controls and each TD group by the independent t test. The results showed all TD groups demonstrated altered correlation patterns of the thumb relative to each finger. Larger forces and higher contributions of the index finger were found during holding by patients with index finger involved, and also during drinking by patients with affected thumb and with affected middle finger. Although no triggering symptom occurred during grasping, the patients showed altered force patterns which may be related to the role of the affected digit in natural grasping function. In conclusion, even if only one digit was affected, the subtle force coordination of all the digits was altered during simple tasks among the TD patients. This study provides the information for the future studies to further comprehend the possible injuries secondary to the altered finger coordination and also to adopt suitable treatment strategies.     

Development and regeneration of the neonatal digit tip in mice

The digit tips of children and rodents are known to regenerate following amputation. The skeletal structure that regenerates is the distal region of the terminal phalangeal bone that is associated with the nail organ. The terminal phalanx forms late in gestation by endochondral ossification and continues to elongate until sexual maturity (8 weeks of age). Postnatal elongation at its distal end occurs by appositional ossification, i.e. direct ossification on the surface of the terminal phalanx, whereas proximal elongation results from an endochondral growth plate. Amputation through the middle of the terminal phalanx regenerates whereas regenerative failure is observed following amputation to remove the distal 2/3 of the bone. Regeneration is characterized by the formation of a blastema of proliferating cells that appear undifferentiated and express Bmp4. Using chondrogenic and osteogenic markers we show that redifferentiation does not occur by endochondral ossification but by the direct ossification of blastema cells that form the rudiment of the digit tip. Once formed the rudiment elongates by appositional ossification in parallel with unamputated control digits. Regenerated digits are consistently shorter than unamputated control digits. Finally, we present a case study of a child who suffered an amputation injury at a proximal level of the terminal phalanx, but failed to regenerate despite conservative treatment and the presence of the nail organ. These clinical and experimental findings expand on previously published observations and initiate a molecular assessment of a mammalian regeneration model.     

Monitoring changes in skin temperature associated with exercise in horses on a water treadmill by use of infrared thermography

Infrared thermography (IRT) was used to assess surface temperature change as an indirect measure of muscle activity and exercise associated changes in blood flow in the working hind limb muscles of horses (n=7) undergoing water treadmill exercise. Three treatments were investigated including the treadmill ran dry (TD), water at the height of the proximal interphalangeal joint (PIP) and water at the height of the carpus (CP). Maximum skin surface temperature was recorded from the region of the semitendinosus muscle during exercise at each water height. There was a significant difference in surface hind limb temperature between exercise on the water treadmill ran dry and with water at the height of the PIP and CP (P<0.0001) with hotter temperatures recorded during the TD treatment. There was a greater increase in surface temperature of the hind limbs from pre exercise to maximum temperature during the PIP and CP treatments when compared to the TD treatment, however, this was not significant (P=0.58). There was no significant difference in surface hind limb temperature found between exercise in water at the height of the PIP and water at the height of the CP. The findings from this study suggest that IRT is able to non-invasively detect muscle activity and associated changes in blood flow whilst horses are exercised on a water treadmill. IRT could potentially be used as an alternative method to assess muscle activity and temperature change in an aquatic environment where existing methods present methodological challenges.     

The Force Synergy of Human Digits in Static and Dynamic Cylindrical Grasps

This study explores the force synergy of human digits in both static and dynamic cylindrical grasping conditions. The patterns of digit force distribution, error compensation, and the relationships among digit forces are examined to quantify the synergetic patterns and coordination of multi-finger movements. This study recruited 24 healthy participants to perform cylindrical grasps using a glass simulator under normal grasping and one-finger restricted conditions. Parameters such as the grasping force, patterns of digit force distribution, and the force coefficient of variation are determined. Correlation coefficients and principal component analysis (PCA) are used to estimate the synergy strength under the dynamic grasping condition. Specific distribution patterns of digit forces are identified for various conditions. The compensation of adjacent fingers for the force in the normal direction of an absent finger agrees with the principle of error compensation. For digit forces in anti-gravity directions, the distribution patterns vary significantly by participant. The forces exerted by the thumb are closely related to those exerted by other fingers under all conditions. The index-middle and middle-ring finger pairs demonstrate a significant relationship. The PCA results show that the normal forces of digits are highly coordinated. This study reveals that normal force synergy exists under both static and dynamic cylindrical grasping conditions.     

Bambi and Sp8 Expression Mark Digit Tips and Their Absence Shows That Chick Wing Digits 2 and 3 Are Truncated

An often overlooked aspect of digit development is the special nature of the terminal phalanx, a specialized structure with characteristics distinct from other phalanges, for example the presence of ectodermal derivatives such as nails and claws. Here, we describe the unique ossification pattern of distal phalanges and characteristic gene expression in the digit tips of chick and duck embryos. Our results show that the distal phalanx of chick wing digit 1 is a genuine tip with a characteristic ossification pattern and expression of Bambi and Sp8; however, the terminal phalanx of digits 2* and 3 is not a genuine tip, and these are therefore truncated digits. Bambi and Sp8 expression in the chick wing provides a direct molecular assessment of digit identity changes after experimental manipulations of digit primordia. In contrast, digits 1 and 2 of the duck wing both possess true tips. Although chick wing-tip development was not rescued by application of Fgf8, this treatment induced the development of extra phalanges. Grafting experiments show that competence for tip formation, including nails, is latent in the interdigital tissue. Our results deepen understanding of the mechanisms of digit tip formation, highlighting its developmental autonomy and modular nature, with implications for digit reduction or loss during evolution. * Numbering of wing digits is 1, 2, 3 from anterior to posterior.     

Comparing the use of live trees and deadwood for larval foraging by aye-ayes (<Emphasis Type="Italic">Daubentonia madagascariensis</Emphasis>) at Kianjavato and Torotorofotsy,Madagascar

Aye-aye (Daubentonia madagascariensis) feeding behavior has become synonymous with deadwood foraging. However, deadwood is not always the most frequently used substrate, as some aye-ayes use live trees more often to access invertebrates. We sought to compare the frequency of aye-aye invertebrate foraging in deadwood and live trees to better understand their feeding behaviors. We followed two male aye-ayes at Kianjavato, a heavily disturbed habitat in southeastern Madagascar, from October 2013 to October 2014, and one male and one female aye-aye at Torotorofotsy, a continuous forest in eastern Madagascar, from July 2014 to December 2015. We collected feeding data by recording the behavior of a focal aye-aye every 5 min for a total of 373 h at Kianjavato and 383 h at Torotorofotsy. Our results showed no difference in the amount of deadwood used between the individuals. However, there was a significant difference in the amount of live tree feeding between the female at Torotorofotsy and one of the males at Kianjavato. We conclude that feeding on invertebrates in live trees is more important to aye-ayes than previously realized and that aye-ayes are exceedingly flexible in their invertebrate feeding behaviors, adjusting to their habitat by using various substrates.     

Even-toed but uneven in length: the digits of artiodactyls

In captive ruminants housed in small enclosures, hypertrophy of the outer hooves of the hindlimbs is often observed. We hypothesised that the underlying cause is overload attributable to an asymmetry of the digits, especially with respect to their length. To test this hypothesis, the bones of the digits of four species of artiodactyls, which included 11 wild chamois (Rupicapra rupicapra), 11 captive fallow deer (Dama dama), 11 captive bison (Bison bison) and 11 European moose (Alces alces; 9 wild, 2 captive), were radiographed post mortem and measured using a computer programme. In addition, the dimensions of the outer and inner hooves were measured directly with a calliper. The mean lengths of the epiphysis of the fourth metacarpal/metatarsal bone and the first and second phalanges of the fourth digit were greater than that of the third digit, whereas the third phalanx of the third digit had a greater mean length. The mean total length of the fourth digit of the forelimbs was greater than that of the third digit in 73–95% of specimens, depending on species. In the hindlimbs, the fourth digit was longer in 91–100% of the specimens. The hooves of the fourth digit were significantly broader than the hooves of the third digit, whereas the inner hooves of the third digits had a greater toe length than those of the fourth digit. The paired digits of artiodactyls are uneven in length, which suggests a different function during stance and weight bearing. It is conceivable that this asymmetry is the result of selection processes that favoured locomotion on soft ground.     

DATA AND DATA INTERPRETATION IN THE STUDY OF LIMB EVOLUTION: A REPLY TO GALIS ET AL. ON THE REEVOLUTION OF DIGITS IN THE LIZARD GENUS BACHIA

Galis and collaborators (2010) claim that our recent paper ( Kohlsdorf and Wagner 2006 ), presenting statistical evidence for the reevolution of digits in the genus Bachia, may be flawed. Their reanalysis of the data does not support the possibility of a reevolution of digits and the authors also argue that such a reevolution would be implausible on functional and developmental grounds. In response, we reanalyzed our data with additional outgroup species. Our results differ from the one published in 2006, but this incongruence is not statistically significant. In contrast, the hypothesis presented by Galis et al. is significantly worse. An analysis of digit number evolution, using novel techniques to test for irreversibility of character loss ( Goldberg and Igic 2008 ), confirmed our original conclusion that there is strong evidence for reevolution of digits in Bachia. We also point out that this result is not in conflict with the hypothesis by Galis and Metz (2001) that mutations affecting the initial digit patterning are associated with strong negative pleiotropic effects and thus unlikely to be fixed in evolution. An important avenue of future research will be to directly test whether reevolved digits develop from conserved digit condensations retained after digit loss.     

A tendon-locking mechanism in two climbing rodents,Muscardinus avellanarius and Micromys minutus (Mammalia,Rodentia)

Having compared the microanatomy of the toes of a terrestrial to two climbing species, adaptations were found in the flexor tendons and in the integument. In contrast to Crocidura russula, both Muscardinus avellanarius and Micromys minutus have a tendon-locking mechanism (TLM) that is engaged when the middle phalanx is bent. A ventral thickening of the flexor tendon is situated deep to a thickened portion of the ventral tendon sheath. When twigs or stalks are grasped, the TLM allows less muscular energy to be expended. In C. russula glands are restricted to the terminal pads, but in the climbing species they occur in the sole of the toes as well. In the reed-living M. minutus knob-shaped integumental thickenings, together with the digital pads, stabilize the grip. In contrast the arboreal M. avellanarius often climbs thick branches and shows adaptations for pressing the sole of the feet against the surface. Thereby the tendon attached to the plantar integument of the toes transfers the muscle force directly to the bark. Unlike the other digits on the forefeet of both climbing species, no TLM is present in the anterior digit. In M. minutus this short digit is twisted towards the palm and, with the carpal pads, provides an abutment against the grasping fingers. In M. avellanarius the anterior digit has very thin tendons and is that much reduced in length that it is completely integrated into the digital pad where it acts, at best, as a lateral support of the pad. © 1996 Wiley-Liss, Inc.     

Hand and body position during locomotor behavior in the aye-aye (Daubentonia madagascariensis)

Aye-ayes (Daubentonia madagascariensis) have unique hands among primates, with extraordinarily long fingers in relation to body size. These long digits may be vulnerable to damage from forces during locomotion, particularly during head-first descent-a locomotor mode that the aye-aye utilizes frequently. Previous behavioral studies of aye-aye locomotion reported that Daubentonia must curl its fingers during horizontal quadrupedalism and/or descent to reduce potential stresses on its long fingers. To test this hypothesis, we examined hand and body position in three captive adult aye-ayes while they walked quadrupedally on horizontal and oblique branches. Substantial variation in hand position was observed among individuals for each substrate orientation. While hand postures with curled fingers were preferred by one individual during descent, they were not preferred by the other two individuals, contrary to our expectations. Differences in body position were more consistent among all three individuals. The angle of the body relative to the substrate was significantly reduced during descent (8.4 degrees ) compared to horizontal locomotion (16.9 degrees ). These results suggest that changes in body position, rather than hand position, may help reduce stresses on the digits. A biomechanical model is proposed that demonstrates how a reduction in the body angle in relation to substrate may act to move the center of mass more caudally. This mechanism of moderating loads by altering body position, rather than hand position, may represent an important functional aspect of arboreal locomotion in aye-ayes and other primates.     

Wear Resistance and Non-Magnetic Layer Formation on 316L Implant Material with Plasma Nitriding

In this study, the applicability of plasma nitriding treatment in the production of non-magnetic and corrosion resistant layer on 316L stainless steel implant material was investigated. 316L stainless steel substrates were plasma nitrided at temperatures of 350 ℃, 375 ℃, 400 ℃, 425 ℃ and 450 ℃ for 2 h in a gas mixture of 50% N2-50% H2, respectively. It was determined that the treatment temperature is the most important factor on the properties of the corrosion resistant layer of 316L stainless steel. The results show that s-phase formed at the temperatures under 400 ℃, and at the temperatures above 400 ℃, instead of s-phase, CrN and y＇-Fe4N phases were observed in the modified layer. The electrical resistivity and surface roughness of the modified layer increase with treatment temperature. Under 400 ℃ the corrosion resistance increased with the temperature, above 400 ℃ it decreased with the increase in treatment temperature. It was analyzed that the electrical resistivity and the soft （ideal） ferro- magnetic properties of 316L stainless steel increased with treatment temperature during nitriding treatment. Also, plasma ni- triding at low temperatures provided magnetic behavior close to the ideal untreated 316L stainless steel.     

Movements of Individual Digits in Bimanual Prehension Are Coupled into a Grasping Component

The classic understanding of prehension is that of coordinated reaching and grasping. An alternative view is that the grasping in prehension emerges from independently controlled individual digit movements (the double-pointing model). The current study tested this latter model in bimanual prehension: participants had to grasp an object between their two index fingers. Right after the start of the movement, the future end position of one of the digits was perturbed. The perturbations resulted in expected changes in the kinematics of the perturbed digit but also in adjusted kinematics in the unperturbed digit. The latter effects showed up when the end position of the right index finger was perturbed, but not when the end position of the left index finger was perturbed. Because the absence of a coupling between the digits is the core assumption of the double-pointing model, finding any perturbation effects challenges this account of prehension; the double-pointing model predicts that the unperturbed digit would be unaffected by the perturbation. The authors conclude that the movement of the digits in prehension is coupled into a grasping component.     

Ectopic dermal ridge configurations on the interdigital webbings and postaxial marginal portion of the hindlimb in Hammertoe mutant mice (Hm)

The effects of the hereditary malformation of Hammertoe mutant mice (gene symbol Hm) on the digital pads and dermal ridge configurations on their hindlimbs were examined. In the wild‐type (+/+) mice with normally separated digits, dermal ridges developed only on the pads. Heterozygous (Hm/+) and homozygous (Hm/Hm) mutant mice, however, had a broad big toe, fused interdigital soft tissues, reduced claws, an extra rudimentary postaxial digit and camptodactyly. The dermal ridges appeared not only on the pads, affected in their number and configurations, but also on the ventral surface of the interdigital webbings and postaxial marginal area exhibiting an extra rudimentary digit and webbing. These aberrant configurations may be related to the abnormal occurrence of programmed cell death (PCD) in the interdigital zones and the postaxial marginal portion in Hm/+ and Hm/Hm mice. That is, the diminished cell death may fail to decrease the cell density in the interdigital zones and postaxial marginal portion and result in the webbing and an extra rudimentary digit and webbing, respectively. Simultaneously, it could also interrupt the migration of surviving cells of these areas toward the neighboring digits and the distal area of the sole and produce the ectopic dermal ridges on the way to the as yet unformed (presumptive) digital and plantar volar pads. The present findings suggest that normal interdigital and pre/postaxial PCD contributes not only to the separation of digits, the initial formation of individual digits of different sizes, and the inhibition of the extra digit but also to the development of the presumptive digital and plantar pads, including dermal ridges. J. Morphol., 2008. © 2008 Wiley‐Liss, Inc.     

Afferent Modulation of Warmth Sensation and Heat Pain in the Human Hand

The hands of 14 normal humans were used to determine the somatotopic organization of the modulation of warmth sensation and heat pain by different forms of cutaneous stimuli. Test stimuli were 5-sec heat pulses ranging from 36° to 51°C, delivered to the fingerpads of digits 1, 2, 4, and 5 with a contact thermode. Conditioning stimuli (15 sec) bracketed the test stimuli and included vibration, noxious and innocuous heat, cold, and electrical pulses delivered to the fingerpads of digits that were adjacent or nonadjacent to the tested digits. Noxious (48° ± 1.3°C), but not innocuous (43°C), heat stimuli increased the perceived magnitude estimation of innocuous test stimuli (36–43°C) by 20–37% when delivered to adjacent, but not to nonadjacent, digits. No other conditioning stimuli had any effect on the intensity of warmth perception. In contrast, both noxious and innocuous heat or electrical conditioning reduced the magnitude estimation of noxious (50–5°C), but not innocuous, test pulses by 12–22% when delivered to adjacent digits. Conditioning of nonadjacent digits was significantly less effective. The analgesic effects of noxious and innocuous conditioning were approximately equal. Vibratory (120 Hz, 3.5 μm) and cold (15°C) conditioning stimuli were ineffective. The results are consistent with a dermatomal somatotopic organization of tactile and heat modulatory influences on warmth sensation and heat pain. The results further suggest that the neural mechanisms subserving warmth mediate a negative feedback influence on heat pain intensity.