Forelimb

Last updated
Forelimbs in mammals have varying functions but are all homologous. Handskelett MK1888.png
Forelimbs in mammals have varying functions but are all homologous.

A forelimb or front limb is one of the paired articulated appendages (limbs) attached on the cranial (anterior) end of a terrestrial tetrapod vertebrate's torso. With reference to quadrupeds, the term foreleg or front leg is often used instead. In bipedal animals with an upright posture (e.g. humans and some primates), the term upper limb is often used.

Contents

A forelimb is not to be confused with a forearm, which is a distal portion of the human upper limb between the elbow and the wrist.

All vertebrate forelimbs are homologous, meaning that they all evolved from the same structures. For example, the flipper of a turtle or of a dolphin, the arm of a human, the foreleg of a horse, and the wings of both bats and birds are ultimately homologous, despite the large differences between them. [1]

Specific uses of the forelimbs may be analogous if they evolved from different sub-structures of the forelimb, such as the flippers of turtles and dolphins, and the wings of birds and bats. [2]

Evolution of forelimbs

Evolution of the forelimb may be characterized by many trends. The number of digits, their characteristics, as well as the shape and alignment of radius, ulna, and humerus, have had major evolutionary implications.

Changes in body size, foot posture, habitat, and substrate are frequently found to influence one another (and to connect to broader potential drivers, such as changing climate). [3]

Shape

A number of factors can influence the evolution of forelimb long bone shape, such as body mass, lifestyle, predatory behavior, or relative prey size. A general pattern is for heavier species to have more robust radii, ulnas, and humeri. [4]

Musteloid carnivorans that have an arboreal lifestyle tend to have long and slender forelimb long bones, which allow for improved movement and flexibility. Semi-fossorial and aquatic musteloid species tend to have short and robust forelimb long bones to deal with the strain from digging and swimming. [5]

In the order Carnivora, felids, which usually ambush and grapple with their prey, have shorter and more robust limbs. Their forelimbs are used for both short sprints and grappling, which means that they need to be flexible and durable. In contrast, canids, which often pursue their prey over greater distances, have longer, more gracile limbs. Running is pretty much the only use for their forelimbs, so they do not need to be adapted for anything else and can be less flexible. [6]

Predators hunting prey that is half their body weight or greater evolved shorter and more sturdy radii, ulnas, and humeri to decrease the likelihood of the bone breaking or fracturing while hunting. Predators hunting prey less than half their body weight tended to have longer and more slender forelimb long bones to improve energetic efficiency. [7]

Polydactyly

Tetrapods were initially understood to have first developed five digits as an ancestral characteristic, which were then reduced or specialized into a number of uses. Certain animals retained 'primitive' forelimbs, such as pentadactylous (five-fingered) reptiles and primates. This has mostly held true, but the earliest tetrapod or "fishapod" ancestors may have had more than five digits. This was notably challenged by Stephen Jay Gould in his 1991 essay "Eight (Or Fewer) Little Piggies". [8]

Polydactyly in early tetrapods should be understood as having more than five digits to the finger or foot, a condition that was the natural state of affairs in the very first tetrapods. Early groups like Acanthostega had eight digits, while the more derived Ichthyostega had seven digits, the yet-more derived Tulerpeton had six toes.

Tetrapods evolved from animals with fins such as found in lobe-finned fishes. From this condition a new pattern of limb formation evolved, where the development axis of the limb rotated to sprout secondary axes along the lower margin, giving rise to a variable number of very stout skeletal supports for a paddle-like foot.

Digit specialization

Digits may be specialized for different forms of locomotion. A classic example is the horse's development of a single toe (monodactyly). [3] Other hooves, like those of even-toed and odd-toed ungulates, and even the hoof-like foot of extinct hadrosaurs, [9] may be regarded as similar specializations.

To bear their immense weight, sauropods, the most derived being titanosaurs, developed a tubular manus (front foot) and gradually lost their digits, standing on their metacarpals. [10] The stegosaurian forelimb has evidence for a sauropod−like metacarpal configuration [11] This was a different evolutionary strategy than megafaunal mammals such as modern elephants.

Therapsids started evolving diverse and specialized forelimbs 270 million years ago, during the Permian. [12]

Opposable thumbs

Modern humans are unique in the musculature of the forearm and hand, though opposable thumbs or structures like them have arisen in a few animals.

In dinosaurs, a primitive autonomization of the first carpometacarpal joint (CMC) may have occurred. In primates, a real differentiation appeared perhaps 70 mya, while the shape of the human thumb CMC finally appears about 5 mya.

Pandas have evolved pseudo-opposable thumbs by extension of the sesamoid bone, which is not a true digit. [14]

Pronation and supination

The ability to pronate the manus (hand) and forearm in therian mammals is achieved by a rounded head of the radius, which allows it to swivel across the ulna. Supination requires a dorsal glide of the distal radius and pronation a palmar glide in relation to the distal ulna.

Pronation has evolved multiple times, among mammals, chameleons, and varanids. [15] However, the more basal condition is to be unable to pronate. Dinosaurs were not capable of more than semi-pronation of the wrist, [16] though bipedal origins of all quadrupedal dinosaur clades could have allowed for greater disparity in forelimb posture than often considered. [15] Monotremes have forearms that are not as dexterous as therians. Monotremes have a sprawling posture, and multiple elements in their pectoral girdles, which are ancestral traits for mammals. [17]

In birds, the forearm muscles supinate, pronate, flex and extend the distal wing. [18]

Wings

All tetrapod forelimbs are homologous, evolving from the same initial structures in lobe-finned fish. However, another distinct process may be identified, convergent evolution, by which the wings of birds, bats, and extinct pterosaurs evolved the same purpose in drastically different ways. [2] These structures have similar form or function but were not present in the last common ancestor of those groups.

Bat wings are composed largely of a thin membrane of skin supported on the five fingers, whereas bird wings are composed largely of feathers supported on much reduced fingers, with finger 2 supporting the alula and finger 4 the primary feathers of the wing; there are only distant homologies between birds and bats, with much closer homologies between any pair of bird species, or any pair of bat species.

Flippers

Marine mammals have evolved several times. Over the course of their evolution, they develop streamlined hydrodynamic bodies. The forelimb thus develops into a flipper. The forelimbs of cetaceans, pinnipeds, and sirenians presents a classic example of convergent evolution. There is widespread convergence at the gene level. [19] Distinct substitutions in common genes created various aquatic adaptations, most of which constitute parallel evolution because the substitutions in question are not unique to those animals. [20]

When comparing cetaceans to pinnipeds to sirenians, 133 parallel amino acid substitutions occur. Comparing and contrasting cetaceans-pinnipeds, cetaceans-sirenians, and pinnipeds-sirenians, 2,351, 7,684, and 2,579 substitutions occur, respectively. [20]

See also

Bibliography

    Related Research Articles

    <span class="mw-page-title-main">Bipedalism</span> Terrestrial locomotion using two limbs

    Bipedalism is a form of terrestrial locomotion where a tetrapod moves by means of its two rear limbs or legs. An animal or machine that usually moves in a bipedal manner is known as a biped, meaning 'two feet'. Types of bipedal movement include walking or running and hopping.

    <span class="mw-page-title-main">Quadrupedalism</span> Form of locomotion using four limbs

    Quadrupedalism is a form of locomotion where four limbs are used to bear weight and move around. An animal or machine that usually maintains a four-legged posture and moves using all four limbs is said to be a quadruped. Quadruped animals are found among both vertebrates and invertebrates.

    <span class="mw-page-title-main">Tetrapod</span> Superclass of the first four-limbed vertebrates and their descendants

    A tetrapod is any four-limbed vertebrate animal of the superclass Tetrapoda. Tetrapods include all extant and extinct amphibians and amniotes, with the latter in turn evolving into two major clades, the sauropsids and synapsids. Some tetrapods such as snakes, legless lizards, and caecilians had evolved to become limbless via mutations of the Hox gene, although some do still have a pair of vestigial spurs that are remnants of the hindlimbs.

    <span class="mw-page-title-main">Homology (biology)</span> Shared ancestry between a pair of structures or genes in different taxa

    In biology, homology is similarity due to shared ancestry between a pair of structures or genes in different taxa. A common example of homologous structures is the forelimbs of vertebrates, where the wings of bats and birds, the arms of primates, the front flippers of whales, and the forelegs of four-legged vertebrates like dogs and crocodiles are all derived from the same ancestral tetrapod structure. Evolutionary biology explains homologous structures adapted to different purposes as the result of descent with modification from a common ancestor. The term was first applied to biology in a non-evolutionary context by the anatomist Richard Owen in 1843. Homology was later explained by Charles Darwin's theory of evolution in 1859, but had been observed before this, from Aristotle onwards, and it was explicitly analysed by Pierre Belon in 1555.

    <span class="mw-page-title-main">Thumb</span> First digit of the hand

    The thumb is the first digit of the hand, next to the index finger. When a person is standing in the medical anatomical position, the thumb is the outermost digit. The Medical Latin English noun for thumb is pollex, and the corresponding adjective for thumb is pollical.

    <span class="mw-page-title-main">Dactyly</span> Arrangement of digits on hands and feet

    In biology, dactyly is the arrangement of digits on the hands, feet, or sometimes wings of a tetrapod animal. It comes from the Greek word δακτυλος = "finger".

    <span class="mw-page-title-main">Theropoda</span> Clade of dinosaurs

    Theropoda, whose members are known as theropods, is a dinosaur clade that is characterized by hollow bones and three toes and claws on each limb. Theropods are generally classed as a group of saurischian dinosaurs. They were ancestrally carnivorous, although a number of theropod groups evolved to become herbivores and omnivores. Theropods first appeared during the Carnian age of the late Triassic period 231.4 million years ago (Ma) and included the majority of large terrestrial carnivores from the Early Jurassic until at least the close of the Cretaceous, about 66 Ma. In the Jurassic, birds evolved from small specialized coelurosaurian theropods, and are today represented by about 10,500 living species.

    <span class="mw-page-title-main">Upper limb</span> Consists of the arm, forearm, and hand

    The upper limbs or upper extremities are the forelimbs of an upright-postured tetrapod vertebrate, extending from the scapulae and clavicles down to and including the digits, including all the musculatures and ligaments involved with the shoulder, elbow, wrist and knuckle joints. In humans, each upper limb is divided into the arm, forearm and hand, and is primarily used for climbing, lifting and manipulating objects.

    <span class="mw-page-title-main">Flipper (anatomy)</span> Flattened limb adapted for propulsion and maneuvering in water

    A flipper is a broad, flattened limb adapted for aquatic locomotion. It refers to the fully webbed, swimming appendages of aquatic vertebrates that are not fish.

    <span class="mw-page-title-main">Patagium</span> Membranous structure that assists an animal in gliding or flight

    The patagium is a membranous body part that assists an animal in obtaining lift when gliding or flying. The structure is found in extant and extinct groups of flying and gliding animals including bats, birds, some dromaeosaurs, pterosaurs, gliding mammals, some flying lizards, and flying frogs. The patagium that stretches between an animal's hind limbs is called the uropatagium or the interfemoral membrane.

    <span class="mw-page-title-main">Cursorial</span>

    A cursorial organism is one that is adapted specifically to run. An animal can be considered cursorial if it has the ability to run fast or if it can keep a constant speed for a long distance. "Cursorial" is often used to categorize a certain locomotor mode, which is helpful for biologists who examine behaviors of different animals and the way they move in their environment. Cursorial adaptations can be identified by morphological characteristics, physiological characteristics, maximum speed, and how often running is used in life. There is much debate over how to define a cursorial animal specifically. The most accepted definitions include that a cursorial organism could be considered adapted to long-distance running at high speeds or has the ability to accelerate quickly over short distances. Among vertebrates, animals under 1 kg of mass are rarely considered cursorial, and cursorial behaviors and morphology are thought to only occur at relatively large body masses in mammals. There are a few mammals that have been termed "micro-cursors" that are less than 1 kg in mass and have the ability to run faster than other small animals of similar sizes.

    <span class="mw-page-title-main">Iguanodontidae</span> Extinct family of dinosaurs

    Iguanodontidae is a family of iguanodontians belonging to Styracosterna, a derived clade within Ankylopollexia.

    Many vertebrates are limbless, limb-reduced, or apodous, with a body plan consisting of a head and vertebral column, but no adjoining limbs such as legs or fins. Jawless fish are limbless but may have preceded the evolution of vertebrate limbs, whereas numerous reptile and amphibian lineages – and some eels and eel-like fish – independently lost their limbs. Larval amphibians, tadpoles, are also often limbless. No mammals or birds are limbless, but some feature partial limb-loss or limb reduction.

    A facultative biped is an animal that is capable of walking or running on two legs (bipedal), as a response to exceptional circumstances (facultative), while normally walking or running on four limbs or more. In contrast, obligate bipedalism is where walking or running on two legs is the primary method of locomotion. Facultative bipedalism has been observed in several families of lizards and multiple species of primates, including sifakas, capuchin monkeys, baboons, gibbons, gorillas, bonobos and chimpanzees. Different facultatively bipedal species employ different types of bipedalism corresponding to the varying reasons they have for engaging in facultative bipedalism. In primates, bipedalism is often associated with food gathering and transport. In lizards, it has been debated whether bipedal locomotion is an advantage for speed and energy conservation or whether it is governed solely by the mechanics of the acceleration and lizard's center of mass. Facultative bipedalism is often divided into high-speed (lizards) and low-speed (gibbons), but some species cannot be easily categorized into one of these two. Facultative bipedalism has also been observed in cockroaches and some desert rodents.

    A limb is a jointed, muscled appendage of a tetrapod vertebrate animal used for weight-bearing, terrestrial locomotion and physical interaction with other objects. The distalmost portion of a limb is known as its extremity. The limbs' bony endoskeleton, known as the appendicular skeleton, is homologous among all tetrapods, who use their limbs for walking, running and jumping, swimming, climbing, grasping, touching and striking.

    A hindlimb or back limb is one of the paired articulated appendages (limbs) attached on the caudal (posterior) end of a terrestrial tetrapod vertebrate's torso. With reference to quadrupeds, the term hindleg or back leg is often used instead. In bipedal animals with an upright posture, the term lower limb is often used.

    <span class="mw-page-title-main">Matthew Bonnan</span> American paleontologist, professor, singer/songwriter

    Matthew Bonnan is an American paleobiologist, a Professor of Biological Sciences at Stockton University, and as of 2021 a singer/songwriter. His research combines traditional descriptive and anatomical study with computer-aided morphometric analysis and modeling of vertebrate skeletons, and he is the co-discoverer of three new species of dinosaurs. He is the author of the book The Bare Bones: An Unconventional Evolutionary History of the Skeleton, designed to introduce undergraduates and curious lay readers to the anatomy and evolution of the vertebrate skeleton. Bonnan has a music/art outreach project, Once Upon Deep Time, a pop/rock song cycle about the evolution of hearing and our connection to the tree of life.

    <span class="mw-page-title-main">Hand</span> Extremity at the end of an arm or forelimb

    A hand is a prehensile, multi-fingered appendage located at the end of the forearm or forelimb of primates such as humans, chimpanzees, monkeys, and lemurs. A few other vertebrates such as the koala are often described as having "hands" instead of paws on their front limbs. The raccoon is usually described as having "hands" though opposable thumbs are lacking.

    <span class="mw-page-title-main">Bat wing development</span>

    The order Chiroptera, comprising all bats, has evolved the unique mammalian adaptation of flight. Bat wings are modified tetrapod forelimbs. Because bats are mammals, the skeletal structures in their wings are morphologically homologous to the skeletal components found in other tetrapod forelimbs. Through adaptive evolution these structures in bats have undergone many morphological changes, such as webbed digits, elongation of the forelimb, and reduction in bone thickness. Recently, there have been comparative studies of mouse and bat forelimb development to understand the genetic basis of morphological evolution. Consequently, the bat wing is a valuable evo-devo model for studying the evolution of vertebrate limb diversity.

    <i>Lisowicia</i> Genus of giant dicynodont therapsid

    Lisowicia is an extinct genus of giant dicynodont synapsid that lived in what is now Poland during the late Norian or earliest Rhaetian age of the Late Triassic Period, about 210–205 million years ago. Lisowicia is the largest known dicynodont, as well as the largest non-mammalian synapsid, reaching about 4.5 metres (15 ft) long, standing up to 2.6 metres (8.5 ft) tall at the hips and weighing around 5–7 metric tons, comparable in size to modern elephants. It was also one of the last dicynodonts, living shortly before their extinction at the end of the Triassic period. Fossils of a giant dicynodont were known from Poland since 2008, but Lisowicia was not named and officially described as a new species until late 2018.

    References

    1. "Image Gallery: Homo Sapiens. homology: homologies of the forelimb among vertebrates". Encyclopædia Britannica. Retrieved January 27, 2013.
    2. 1 2 "Homologies and analogies". evolution.berkeley.edu. Retrieved 2019-12-09.
    3. 1 2 McHorse, Brianna K.; Biewener, Andrew A.; Pierce, Stephanie E. (2019-09-01). "The Evolution of a Single Toe in Horses: Causes, Consequences, and the Way Forward". Integrative and Comparative Biology. 59 (3): 638–655. doi: 10.1093/icb/icz050 . ISSN   1540-7063. PMID   31127281.
    4. Fabre, Anne-Claire; Cornette, Raphael; Peigné, Stéphane; Goswami, Anjali (2013-05-21). "Influence of body mass on the shape of forelimb in musteloid carnivorans". Biological Journal of the Linnean Society. 110 (1): 91–103. doi: 10.1111/bij.12103 . ISSN   0024-4066.
    5. Fabre, Anne-Claire; Cornette, Raphael; Goswami, Anjali; Peigné, Stéphane (2015-05-21). "Do constraints associated with the locomotor habitat drive the evolution of forelimb shape? A case study in musteloid carnivorans". Journal of Anatomy. 226 (6): 596–610. doi: 10.1111/joa.12315 . ISSN   0021-8782. PMC   4450962 . PMID   25994128.
    6. Meachen-Samuels, Julie; Van Valkenburgh, Blaire (June 2009). "Forelimb indicators of prey-size preference in the Felidae". Journal of Morphology. 270 (6): 729–744. doi:10.1002/jmor.10712. ISSN   0362-2525. PMID   19123240. S2CID   20732642.
    7. Michaud, Margot; Veron, Géraldine; Fabre, Anne‐Claire (2020-11-06). "Phenotypic integration in feliform carnivores: Covariation patterns and disparity in hypercarnivores versus generalists". Evolution. 74 (12): 2681–2702. doi:10.1111/evo.14112. ISSN   0014-3820. PMID   33085081. S2CID   224824184.
    8. Stephen Jay Gould. "Stephen Jay Gould "Eight (or Fewer) Little Piggies" 1991". Archived from the original on 2010-01-11. Retrieved 2015-10-02.
    9. Zheng, R. ; Farke (2011). "A Photographic Atlas of the Pes from a Hadrosaurine Hadrosaurid Dinosaur". PalArch's Journal of Vertebrate Palaeontology. 8 (7): 1–12. ISSN   1567-2158.{{cite journal}}: CS1 maint: multiple names: authors list (link)
    10. Apesteguía, Sebastián (2005-01-01). "Evolution of the titanosaur metacarpus". Thunder-Lizards: The Sauropodomorph Dinosaurs: 321–345.
    11. Senter, Phil (2010). "Evidence for a Sauropod-Like Metacarpal Configuration in Stegosaurian Dinosaurs". Acta Palaeontologica Polonica. 55 (3): 427–432. doi: 10.4202/app.2009.1105 . ISSN   0567-7920.
    12. "Mammals' unique arms started evolving before the dinosaurs existed". ScienceDaily. Retrieved 2019-12-10.
    13. Ankel-Simons, Friderun. (2007). Primate anatomy : an introduction (3rd ed.). Amsterdam: Elsevier Academic Press. ISBN   978-0-08-046911-9. OCLC   437597677.
    14. Salesa, Manuel J.; Antón, Mauricio; Peigné, Stéphane; Morales, Jorge (2006-01-10). "Evidence of a false thumb in a fossil carnivore clarifies the evolution of pandas". Proceedings of the National Academy of Sciences. 103 (2): 379–382. Bibcode:2006PNAS..103..379S. doi: 10.1073/pnas.0504899102 . ISSN   0027-8424. PMC   1326154 . PMID   16387860.
    15. 1 2 VanBuren, Collin S.; Bonnan, Matthew (2013-09-18). "Forearm Posture and Mobility in Quadrupedal Dinosaurs". PLOS ONE. 8 (9): e74842. Bibcode:2013PLoSO...874842V. doi: 10.1371/journal.pone.0074842 . ISSN   1932-6203. PMC   3776758 . PMID   24058633.
    16. Hutson, Joel D. (2014). "Quadrupedal Dinosaurs did not Evolve Fully Pronated Forearms: New Evidence from the Ulna". Acta Palaeontologica Polonica. 60 (3): 599–610. doi: 10.4202/app.00063.2014 . ISSN   0567-7920.
    17. Hall, Brian Keith, 1941- (2007). Fins into limbs evolution, development, and transformation. University of Chicago Press. OCLC   928978489.{{cite book}}: CS1 maint: multiple names: authors list (link) CS1 maint: numeric names: authors list (link)
    18. Tobalske, Bret W. (2007-09-15). "Biomechanics of bird flight". Journal of Experimental Biology. 210 (18): 3135–3146. doi: 10.1242/jeb.000273 . ISSN   0022-0949. PMID   17766290.
    19. Chikina, Maria; Robinson, Joseph D.; Clark, Nathan L. (2016-09-01). "Hundreds of Genes Experienced Convergent Shifts in Selective Pressure in Marine Mammals". Molecular Biology and Evolution. 33 (9): 2182–2192. doi:10.1093/molbev/msw112. ISSN   0737-4038. PMC   5854031 . PMID   27329977.
    20. 1 2 Zhou, Xuming; Seim, Inge; Gladyshev, Vadim N. (2015-11-09). "Convergent evolution of marine mammals is associated with distinct substitutions in common genes". Scientific Reports. 5 (1): 16550. Bibcode:2015NatSR...516550Z. doi: 10.1038/srep16550 . ISSN   2045-2322. PMC   4637874 . PMID   26549748.