Comparative morphology of the tibial flexor and extensor tendons in insects

Abstract. The relative size, orientation, and degrees of sclerotization of the tibial flexor and extensor tendons are compared in nineteen orders of insects. The sclerotized, independently movable tibial flexor sclerite, known previously only from Alticinae and Carabidae (Coleoptera), is found in some other Coleoptera, Megaloptera, Neuroptera, Hymenoptera and Heteroptera. The Heteroptera also have another small sclerite at the base of the tibial extensor tendon. The tibial flexor sclerite is presumed to provide additional strength and leverage to the flexion of the tibia in certain insect groups; it may also provide protection for the ventral side of the femoro-tibial joint of the leg.     

The independent evolution of the metafemoral spring in Coleoptera

Abstract. The metafemoral spring (jumping organ) was known previously only from all Alticinae (Chrysomelidae), one genus of Bruchidae, and two species of Rhynchaeninae (Curculionidae). Here this spring is reported from three subfamilies, seven tribes and twenty-two genera of Curculionidae (three Rhychaeninae, one Erirhininae (Derelomini), and eighteen Ceutorhynchinae) and also from five genera of Buprestidae (four Agrilinae and one Trachyinae). Jumping in Hexapoda is discussed, specifically in the other jumping Coleopteran families that were examined for the presence of the spring (e.g. Melandryidae, Mordellidae, Scraptiidae, Eucinetidae, Limnichidae, Scirtidae and Anthribidae). The phylogenetic value of the metafemoral spring is still unclear; however, there are indications that it is useful in assessing relationships among weevil tribes. As in Alticinae, Curculionidae and Buprestidae have constant intra-generic spring morphology and inter-generic differences. The spring in Bruchidae, Curculionidae and Buprestidae has a simpler morphology than in the Alticinae, but still possesses the features necessary for jumping (metatibial extension). The metafemoral spring has apparently evolved independently in Alticinae, Bruchidae, Curculionidae and Buprestidae, and is an example of convergent evolution.     

Jumping mechanisms and performance in beetles. II. Weevils (Coleoptera: Curculionidae: Rhamphini)

We describe the kinematics and performance of the natural jump in the weevil Orchestes fagi (Fabricius, 1801) (Coleoptera: Curculionidae) and its jumping apparatus with underlying anatomy and functional morphology. In weevils, jumping is performed by the hind legs and involves the extension of the hind tibia. The principal structural elements of the jumping apparatus are (1) the femoro-tibial joint, (2) the metafemoral extensor tendon, (3) the extensor ligament, (4) the flexor ligament, (5) the tibial flexor sclerite and (6) the extensor and flexor muscles. The kinematic parameters of the jump (from minimum to maximum) are 530–1965 m s^?2 (acceleration), 0.7–2.0 m s^?1 (velocity), 1.5–3.0 ms (time to take-off), 0.3–4.4 μJ (kinetic energy) and 54–200 (g-force). The specific joint power as calculated for the femoro-tibial joint during the jumping movement is 0.97 W g^?1. The full extension of the hind tibia during the jump was reached within up to 1.8–2.5 ms. The kinematic parameters, the specific joint power and the time for the full extension of the hind tibia suggest that the jump is performed via a catapult mechanism with an input of elastic strain energy. A resilin-bearing elastic extensor ligament that connects the extensor tendon and the tibial base is considered to be the structure that accumulates the elastic strain energy for the jump. According to our functional model, the extensor ligament is loaded by the contraction of the extensor muscle, while the co-contraction of the antagonistic extensor and flexor muscles prevents the early extension of the tibia. This is attributable to the leverage factors of the femoro-tibial joint providing a mechanical advantage for the flexor muscles over the extensor muscles in the fully flexed position. The release of the accumulated energy is performed by the rapid relaxation of the flexor muscles resulting in the fast extension of the hind tibia propelling the body into air.     

Elastic energy storage in tendons: mechanical differences related to function and age

We investigated the possibility that tendons that normally experience relatively high stresses and function as springs during locomotion, such as digital flexors, might develop different mechanical properties from those that experience only relatively low stresses, such as digital extensors. At birth the digital flexor and extensor tendons of pigs have identical mechanical properties, exhibiting higher extensibility and mechanical hysteresis and lower elastic modulus, tensile strength, and elastic energy storage capability than adult tendons. With growth and aging these tendons become much stronger, stiffer, less extensible, and more resilient than at birth. Furthermore, these alterations in elastic properties occur to a significantly greater degree in the high-load-bearing flexors than in the low-stress extensors. At maturity the pig digital flexor tendons have twice the tensile strength and elastic modulus but only half the strain energy dissipation of the corresponding extensor tendons. A morphometric analysis of the digital muscles provides an estimate of maximal in vivo tendon stresses and suggests that the muscle-tendon unit of the digital flexor is designed to function as an elastic energy storage element whereas that of the digital extensor is not. Thus the differences in material properties between mature flexor and extensor tendons are correlated with their physiological functions, i.e., the flexor is much better suited to act as an effective biological spring than is the extensor.     

The extensor assembly in two species of opossum,Philander opossum and Didelphis marsupialis

In considering primate and hominoid phylogeny, the fundamental position assigned to opossums is explained partially by the characteristic morphology of their hands and feet. One of the main functional features of the human hand is the ability to make a stabilized arch of the finger. Because the extensor assembly plays a key role in establishing an arched finger, the extensor systems of the digits of both the hands and feet were studied in two species of opossum, Philander opossum and Didelphis marsupialis. In the foot, two extensor tendons join in each toe to form one tendinous plate, which inserts onto the base of the second phalanx. Lumbricals join this plate along the tibial side, and interosseus insertions are found, although a true interosseus wing is lacking. At the proximal interphalangeal level, a terminal tendon takes its origin from this tendinous plate. This terminal tendon is oval in cross-section and contains elastic structures. Oblique bands arise from this terminal tendon and run proximally along the proximal interphalangeal joint inserting onto the base of the first phalanx. There are elastic structures in the flexor tendon on the dorsal side near its site of insertion. In the hand, the main extensor tendons are arranged differently and the interossei contribute substantially to the extensor assembly. Otherwise, the extensor assembly of the hands and feet are quite similar. The function of the so-called paratendinous intravaginal flexors is discussed as are evolutionary aspects of the extensor assembly.     

Comparative study of the characteristics and properties of tendinocytes derived from three tendons in the equine forelimb

The aim of this study was to determine the characteristic differences in tendinocytes derived from tendons in the equine forelimb, superficial digital flexor tendon (SDFT), deep digital flexor tendon (DDFT) and common digital extensor tendon (CDET), in morphology, proliferation, collagen production ability and ability for synthesis of matrix metalloproteinases (MMPs). Significant differences were observed in cell number in vivo. The cellular number was largest in the SDFT and smallest in the CDET. The values of in vitro proliferation ratios and ability for synthesis of collagen and MMPs were largest in the SDFT and smallest in the CDET. Addition of TNFα to culture of all three types of tendinocytes increased the synthesis of both proMMP-9 (except CDET) and collagen and decreased proMMP-13 synthesis and had no effect on proMMP-2 synthesis. Flexor tendons in forelimbs (SDFT and DDFT) restore energy during locomotion and are more easily injured than are extensor tendons. This structural property would cause active ECM and MMPs synthesis. And CDET have very low turnover potential; in the small number of cells, low cellular proliferation, lower ability for synthesis of collagen and MMPs. The isolated tendinocytes provided much information on the characteristics and properties of tendons for the ECM turnover system and responsiveness of tendinocytes to complex inflammatory responses in tendinopathy.     

Tenocyte response to cyclical strain and transforming growth factor beta is dependent upon age and site of origin

The effect of strain and transforming growth factor beta on equine tendon fibroblasts (tenocytes) was assessed in vitro. Tenocytes were isolated from flexor and extensor tendons of horses from foetal to 10 years of age. These cells were cultured until confluent on collagen-coated silicone dishes. Cyclic biaxial strain of 9+/-1% was applied at 0.5 Hz for 24 hours with or without added TGFbeta1 or 3 (10 ng/ml). Proliferation and synthetic responses were dependent on the tendon of origin. Neither strain nor TGFbeta caused flexor tenocytes to proliferate significantly, while strain alone did proliferate extensor tenocytes. TGFbeta, with or without strain, increased the incorporation of [3H]-proline and the production of types I and III collagen and COMP in both cell types, although the effect on COMP production was more marked in flexor tenocytes, perhaps reflecting the higher levels found in this tendon in vivo. Immature flexor tenocytes synthesised more collagen and COMP than those from mature animals, while age had little effect in extensor tenocytes. Our results suggest that tenocytes become differentiated at an early age and present tendon-specific responses.     

Anatomical variations of the extensor tendons to the fingers over the dorsum of the hand: a study of 50 hands and a review of the literature

The extensor tendons to the fingers were studied in dissections of 50 fresh cadaveric hands, and the divisions of the tendons, as well as the communications (juncturae), were analyzed. The pattern of distribution most frequently observed was as follows. The extensor digitorum communis provided one tendon to the index finger, one to the middle finger, two to the ring finger, and none to the little finger. The extensor indicis exhibited one tendon, whereas the extensor digiti minimi exhibited two tendons. The extensor indicis tendon was always observed to lack a junctura tendinum. The extensor indicis was absent in both hands of one cadaver. A tendon slip from the extensor digiti minimi to the ring finger was observed in one hand. All surgeons must bear in mind the existence of these variations when performing common tendon transfers.     

叶甲科三亚科十三种叶甲酯酶同工酶的研究(鞘翅目,叶甲总科)

应用聚丙烯酰胺凝胶电泳方法分析研究了叶甲科3亚科13种昆虫的酯酶同工酶.结果显示,其聚类结果与传统分类结果基本相一致,说明以酯酶同工酶作为研究手段来进行叶甲类昆虫亚科以下阶元的分类是可行的,同时也说明了它们酯酶同工酶酶谱的差异和其分类地位是一致的;但跳甲亚科和叶甲亚科先聚为一类,再与萤叶甲亚科聚为一类,与前人的研究有差异,作者认为:酯酶同工酶的编码基因可能是快进化单位,在解决亚科以下阶元的系统关系时是很好的分子标记,而对于研究叶甲科、亚科间的系统关系,就不一定很合适.     

跳甲的食性及食性分化

跳甲隶属于鞘翅目叶甲科跳甲亚科,是一类具有重要经济意义的植食性昆虫。本文对跳甲食性及食性分化的研究进展进行了综述,诠释了寄主植物的概念,分析了昆虫食性专化发生的原因。跳甲亚科的寄主植物的范围虽广,但有85%的属为专食性属。一般认为,专食性的跳甲亚科和萤叶甲亚科是由食性较广的叶甲亚科进化而来的,食性分化在其中可能起了重要作用。广食性代价推动了食性分化的发生,广食性代价假说受到越来越多的实验支持。有关食性分化方向的观点倾向于寄主植物转向化学物质相似的植物,表现为寄主转移、寄主扩张和形成寄主型等;食性分化推动了同域物种形成。跳甲食性分化的研究对于丰富研究昆虫与植物相互关系的协同进化理论也有重要作用。     

The effect of tendon loading on in-vitro carpal kinematics of the wrist joint

Measurements of in-vitro carpal kinematics of the wrist provide valuable biomechanical data. Tendon loading is often applied during cadaver experiments to simulate natural stabilizing joint compression in the wrist joint. The purpose of this study was to investigate the effect of tendon loading on carpal kinematics in-vitro.A cyclic movement was imposed on 7 cadaveric forearms while the carpal kinematics were acquired by a 4-dimensional rotational X-ray imaging system. The extensor- and flexor tendons were loaded with constant force springs of 50 N, respectively. The measurements were repeated without a load on the tendons. The effect of loading on the kinematics was tested statistically by using a linear mixed model.During flexion and extension, the proximal carpal bones were more extended with tendon loading. The lunate was on the average 2.0° (p=0.012) more extended. With tendon loading the distal carpal bones were more ulnary deviated at each angle of wrist motion. The capitate was on the average 2.4° (p=0.004) more ulnary deviated.During radioulnar deviation, the proximal carpal bones were more radially deviated with the lunate 0.7° more into radial deviation with tendon loading (p<0.001). Conversely, the bones of distal row were more flexed and supinated with the capitate 1.5° more into flexion (p=0.025) and 1.0° more into supination (p=0.011).In conclusion, the application of a constant load onto the flexor and extensor tendons in cadaver experiments has a small but statistically significant effect on the carpal kinematics during flexion–extension and radioulnar deviation.     

蚊蝎蛉捕捉足构造

依据光学显微镜和扫描电镜观察,对扁蚊蝎蛉Bittacus planus Cheng和缠绕蚊蝎蛉Bittacus implicatus Huang et Hua成虫足的外部形态、肌肉组织及超微结构进行了描述和绘图。蚊蝎蛉成虫三对足结构相似,均为捕捉式,以高度特化的跗节捕捉猎物,各跗分节间具发达的关节、凹槽和爪缩肌肌腱膨大区,第4和5跗分节的齿形成嵌合构造,在昆虫纲中为该类群独有特征。首次发现胫节伸肌b由腿节基部发出的一小块三角形肌肉组织和一条长肌腱组成; 爪缩肌肌腱在第5跗分节凹槽末端有一突起,收缩时突起羁绊于凹槽内的“Y”字形底托。分析了成虫捕食行为与捕捉足构造之间的关系,并简要比较了蚊蝎蛉捕捉足与螳螂捕捉足的结构特点。     

The oldest genus of the subfamily Sagrinae (Coleoptera: Chrysomelidae) from the Paleocene of Menat (France)

The new oldest representative of the subfamily Sagrinae, Gallopsis perita gen. et sp. n. from the Paleocene of Menat (France), is described and illustrated. It is distinguished from the genus Neodiaphanops Blackburn, 1899 by the metafemora without teeth, wider pronotum, weakly widened tibiae, bilobed tarsomere 2, subparallel elytra, and narrower metanepisterna. It differs from the Australian genus Carpophagus Macleay, 1826 in the head without rostrum, weakly widened metafemora without teeth, and gently sculptured pronotum and elytra. The new genus differs from the Eocene genus Eosagra Haupt, 1950 in the longer antennomere 2, wider pronotum and elytra, dense and diffuse elytral puncturation, and weakly inflated metafemora.     

Convergent evolution of chemical defence in Galerucine larvae

Selection pressure by natural enemies on phytophagous insect larvae is intense and has frequently triggered the evolution of chemical defence as an effective counterstrategy. In the chrysomelid subfamily Galerucinae, glandular structures and defensive fluids have been described for the tribe Sermylini Wilcox, 1965. Previous morphological and ultrastructural studies raised doubts that these defensive devices in Sermylini can be traced back to a common origin. The taxonomy of the Galerucinae cannot clear these doubts because the phylogeny of this taxon is a matter of current debate. We therefore investigated the phylogeny of the Galerucinae based on approximately 1740 bp of the mitochondrial 12S and 16S rRNA and the nuclear elongation factor 1 alpha genes. Our data support the hypothesized close relationship between the subfamilies Galerucinae and Alticinae, yet, by contrast to other recent analyses, the two groups are mostly resolved as monophyletic sister groups or, in some analyses, with the Galerucinae nested paraphyletically within the Alticinae. Within the subfamily Galerucinae, only the tribe Galerucini formed a monophyletic taxon, except for one species, Cerochroa brachialis Stal, 1858. In none of our analyses were the Sermylini recovered as a monophyletic tribe. However, our data support monophyly of each of the three groups within the Sermylini that have morphologically distinguishable larval defensive openings. We conclude that the defensive structures in larvae of Sermylini have no common origin, but evolved independently. Our data suggest that the tremendous selection pressure by natural enemies led to the recurrent evolution of similar chemical defensive devices in Sermylini larvae.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 93 , 165–175.     

Proprioceptive regulation of locomotion

Recent investigations of proprioreceptors in the walking systems of cats, insects and crustaceans have identified reflex pathways that regulate the timing of the transition from stance to swing, and control the magnitude of ongoing motoneuronal activity. An important finding in the cat is that during locomotor activity, the influence of feedback from the Golgi tendon organs in extensor muscles onto extensor motoneurons is reversed from inhibition to excitation. The excitatory action of tendon organs during stance ensures that stance is maintained while extensor muscles are loaded, and may regulate the magnitude of extensor activity according to the load carried by the leg. Afferents from primary and secondary spindles in extensor and flexor muscles have also been found to influence the timing of the locomotor rhythm in a functionally relevant manner. Recent studies indicate that reflex reversals and the regulation of timing by multiple proprioceptive systems are also features of walking systems in arthropods.     

Organization of connective tissue patterns by dermal fibroblasts in the regenerating axolotl limb

A set of tendons, aponeurotic sheets and retinaculae, which transduce muscle action from proximal limb levels to flexion and extension of the digits, is found in limbs of many vertebrates. This set of structures, here termed the digit tendon complex, is described for the axolotl forelimb. We show that the complex forms autonomously in muscleless axolotl limb regenerates produced from a cuff of unirradiated dermis surrounding an irradiated limb stump, and persists for up to a year after amputation. The pattern of other connective tissue structures, including the skeleton, is also normal. Fibroblast condensations that may represent sets of these cells normally associated with muscles in the extensor and flexor compartments of the carpal region also form in muscleless limbs. The results are discussed in terms of the importance of the dermis in pattern regulation, selforganization of connective tissues in general and autonomous development of the digit tendon complex in particular.     

Neural circuits for jumping in the locust

The locust jump consists of three distinct phases: Cocking: a rapid flexion of both hindleg tibia and locking of both tibia in full flexion. Co-contraction: simultaneous contractions in hindleg flexor and extensor muscles lasting about 0.5 s resulting in the storage of energy for the jump in elastic elements of the legs and muscles. Triggering: a sudden inhibition of flexor activity to allow the shortening of the contracted extensors and the release of the energy stored during the co-contraction phase. The neural circuitry controlling these three phases is now reasonably well understood. Some of its major features are: (1) pairs of large identifiable interneurons in the thoracic ganglia for evoking the cocking response (C-neurons) and for triggering the jump (M-neurons), (2) a central excitatory pathway from extensor to flexor tibiae motoneurons to ensure simultaneous activation of extensor and flexor motoneurons during the initial part of the co-contraction phase, (3) a positive feedback pathway from cuticular receptors to extensor motoneurons for maintaining extensor activity during the co-contraction phase, (4) proprioceptive feedback to the trigger interneurons for increasing their excitability during the co-contraction phase and thereby allowing a variety of external stimuli to activate the trigger neurons and evoke a jump, (5) presynaptic inhibition of visual pathways to the trigger neurons to ensure that the trigger neurons are not activated by the simultaneous occurrence of visual and auditory stimuli in the absence of proprioceptive input, and (6) a pair of multifunctional visual movement detecting neurons which can initiate cocking or trigger the jump depending on the animal's state.     

Macroscopic and microscopic analyses in flexor tendons of the tarsometatarso‐phalangeal joint of ostrich (Struthio camelus) foot with energy storage and shock absorption

Flexor tendons function as energy storage and shock absorption structures in the tarsometatarso‐phalangeal joint (TMTPJ) of ostrich feet during high‐speed and heavy‐load locomotion. In this study, mechanisms underlying the energy storage and shock absorption of three flexor tendons of the third toe were studied using histology and scanning electron microscopy (SEM). Macroscopic and microscopic structures of the flexor tendons in different positions of TMTPJ were analyzed. Histological slices showed collagen fiber bundles of all flexor tendons in the middle TMTPJ were arranged in a linear‐type, but in the proximal and distal TMTPJ, a wavy‐type arrangement was found in the tendon of the M. flexor digitorum longus and tendon of the M. flexor perforans et perforatus digiti III, while no regular‐type was found in the tendon of the M. flexor perforatus digiti III. SEM showed that the collagen fiber bundles of flexor tendons were arranged in a hierarchically staggered way (horizontally linear‐type and vertically linear‐type). Linear‐type and wavy‐type both existed in the proximal TMTPJ for the collagen fiber bundles of the tendon of the M. flexor perforatus digiti III, but only the linear‐type was found in the distal TMTPJ. A number of fibrils were distributed among the collagen fiber bundles, which were likely effective in connection, force transmission and other functions. The morphology and arrangement of collagen fiber bundles were closely related to the tendon functions. We present interpretations of the biological functions in different positions and types of the tendons in the TMTPJ of the ostrich feet.