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Functional histology of quill erection in the porcupine, Erethizon dorsatum
Canadian Journal of Zoology
Abstract
The histology of the porcupine's skin and subcutis is described. The functional relationships of the thoracic skin during elevation of the quills involve a smooth arrector muscle attached to quill sets, one set behind the other, with the muscle at a slant usually from a single root tip of one set to the distal part of the follicles in the caudal set. Stretched elastin returns the quills to their resting position. There is a low-resistance pathway for the tilting quill roots through lease connective tissue and fat until a collagenous strand attached to the quill set stops further tilt. The lumbosacral region (rosette) has a two-layered arrector, but it attaches to only its own quill set and inserts into the dermis. Both regions have a varied and complicated system of transverse and oblique muscles associated with the roots in a set. (The occurrence of obliques is variable in the thoracic region.) A hypothesis is presented concerning the mechanism of quill detachment from the porcupine after the quill is rammed into a predator. This involves the shearing of the conical root tip along its adherent follicular cells and a stopping device (spool) to prevent the root from self-stabbing.
... researchers (Chapman et al. 2018;Cho et al. 2012;Roze 2009Roze , 2012Castilho et al. 2013;Chapman and Roze 1997;Roze 2002;Vincent and Owers 1985). Several behavioral and morphological adaptations associated with porcupine mechanisms of defense have already been described for the North American porcupine, Erethizon dorsatum (Chapman et al. 2018;Cho et al. 2012;Roze 2009Roze , 2012Chapman and Roze 1997;Roze 2002). ...
... researchers (Chapman et al. 2018;Cho et al. 2012;Roze 2009Roze , 2012Castilho et al. 2013;Chapman and Roze 1997;Roze 2002;Vincent and Owers 1985). Several behavioral and morphological adaptations associated with porcupine mechanisms of defense have already been described for the North American porcupine, Erethizon dorsatum (Chapman et al. 2018;Cho et al. 2012;Roze 2009Roze , 2012Chapman and Roze 1997;Roze 2002). In general, the defense repertoire of this species begins with the erection of omnidirectional quills and the presentation of its back to the potential aggressor, exhibiting an apparently larger body size, aposematic color, and warning odor (Roze 2009(Roze , 2012Li, Roze, and Locke 1997). ...
... In general, the defense repertoire of this species begins with the erection of omnidirectional quills and the presentation of its back to the potential aggressor, exhibiting an apparently larger body size, aposematic color, and warning odor (Roze 2009(Roze , 2012Li, Roze, and Locke 1997). Among the morphological characteristics involved in this process are the presence of piloerector muscles, a fluorescent and contrasting color in the quill surface, odor glands, and osmetrichia quills (quills with modifications for odor dispersal) in the dorsum region (Roze 2009(Roze , 2012Chapman and Roze 1997;Li, Roze, and Locke 1997). Erect porcupine quills can penetrate the skin and muscular tissue of an opponent when touched, and the presence of microscopic overlapping, backward-facing barbs in the tip facilitates the penetration (Cho et al. 2012). ...
New World porcupines (Erethizontidae) exhibit behaviors and possess integumentary structures, including the quills, that are used for self‐defense. The North American porcupine ( Erethizon dorsatum ) has been well studied regarding these features; however, information is lacking for the South American Coendou species. We describe the defensive behavior and integumentary morphology of Coendou spinosus to understand the defensive strategies of this species and to compare with those reported for other species. We assessed the behaviors related to warning, defense, and escape of eight porcupines, as well as the characteristics of their pelage and quills. Furthermore, we microscopically analyzed skin samples of a roadkill adult male specimen. Similar to E. dorsatum , C. spinosus exhibited omnidirectional quill erection, revealing an aposematic color and, with their backs toward the perceived human threat, they performed quick tail and body movements to strike the hands of the human trying to capture them by the tail. Furthermore, C. spinosus presented an integumentary structure similar to that of E. dorsatum , and mechanisms to facilitate quill release when touched, penetration, and fixation in the opponent. The most distinct warning behavior noted was the vibration of the quills, which has not been reported for Erethizon . Our study confirms that, like other erethizontids, C. spinosus does not attack but exhibits warning, defense, and escape mechanisms and behaviors when threatened or touched. The dissemination of such information helps to counter the negative stigma associated with porcupines, as they can be the victims of attacks by dogs and humans, and to promote their conservation.
... Deep to the cuticle is the substantial cortex composed of elongated cylindrical keratinized cells bluntly pointed at the ends. In Erethizon, the medullary center is a foam of cell surfaces akin to a mass of polyhedral bubbles; however, many of the finest hairs lack a medulla (Chapman and Roze 1997). ...
... Proximal to the barbed tip, the quill shaft lacks an imbricated cuticle; instead, the cuticular cells are flat tiles. Proceeding basally, the quill narrows drastically to give a slender segment (root or neck), which ends in a minute anchoring conical root tip, which is part of a detachment mechanism that facilitates easy separation from the skin of the porcupine following contact with a predator (Chapman and Roze 1997). ...
... Elastin fibers radiate from this junction (Fig. 1e). Erethizon, on the other hand, has its root tip stabilized by two collagen strands (Chapman and Roze 1997). ...
This study examines the functional histology of the thin-spined porcupine's (Chaetomys subspinosus (Olfers, 1818)) quills and integument, and compares them with the North American porcupine (Erethizon dorsatum (Linnaeus, 1758)) equivalents. Erethizon quills disseminate warnings to potential predators. An olfactory warning is generated by sebaceous glands of the lower back, disseminated by osmetrichial specializations of lower-back quills. Chaetomys skin lacks functional sebaceous glands, and generates no warning odor. Erethizon quills also carry a visual warning, effective even in dim light. Chaetomys quills carry no warning color. The straight, stiff quills of Erethizon are tipped with one-way barbs and can be elevated to 90°. A protective collagenous spool at the base of the quill prevents back-stabbing on impact. Chaetomys quills lack barbs and the basal collagenous spool is poorly developed. The quill shafts are wavy and angled at the base, allowing them to act as shock absorbers during a fall from a tree. Nevertheless, Chaetomys quills of the head and shoulders can be elevated to 90° to function as stabbing devices. Major defensive quills of Erethizon protect the back; in Chaetomys, the front. Though both species have 1–3 arrector muscles, they differ in the details of quill stabilization. The Chaetomys quill surface has regional cuticular patterns, of which the distal, longest region is highly pitted. The function of the pitted region remains a mystery.
... The structure and chemical composition of the skin have affected many physiological and other conditions of the skin such as wound healing (Burd et al., 1990;Cuttle et al., 2005;Liu et al., 1997;Merkel et al., 1988;Volk et al., 2011), bacterial population (Grice et al., 2008) and defence mechanism (Chapman & Roze, 1997;Zherebtsova, 2000). Meanwhile, the skin is composed of two primary layers, namely the epidermis and the dermis. ...
... Histological examination has shown that quill follicles are composed of connective tissue of various densities, an abundance of fat tissue, well-developed sebaceous glands, erector muscles and an epidermal lining (Prawira et al., 2019;Prawira, Hanadhita, et al., 2018;Prawira, Novelina, et al., 2018). The arrangement of quill follicles, connective, erector and subcutaneous muscles in the skin play an important role in quill erection and contribute to the defence mechanism of the porcupine (Chapman & Roze, 1997;Zherebtsova, 2000). ...
... These results were in accordance with the previous study (Prawira, Hanadhita, et al., 2018) which exhibited the thickest quill follicle sheath was found in the dorsal lumbar region where the biggest and hardest quill grow. A study of North American porcupine (Erethizondorsatum) by Chapman and Roze (1997) showed that the collagenous spool covers the base of the quill and serves as a structure that limits the backward movement of the quill into dermis and prevents the quill from selfpenetrating. In addition, when this quill extends vigorously, this structure serves as a fulcrum and stabilized it rigidly. ...
The complexity of the Sunda porcupine skin has become an important topic due to the unique characteristics of its quill follicles. The structure and chemical composition of the skin has affected many physiological and other conditions. Generally, quills are larger, stronger and stiffer than hair; therefore, the skin structure needs to adapt to support their physiology. The strength of the skin is determined by its collagen composition and arrangement; therefore, this study aims to analyse the composition and distribution of thick and thin fibres based on the specific characteristics of Sunda porcupine skin under polarized light using picrosirius red staining. The skin samples used were from the thoracodorsal and lumbosacral regions of eight Sunda porcupine adults. The histological staining was carried out using the picrosirius red method, while the samples were observed under a polarized light microscope and analysed with software. The results showed that the skin is composed of 36%–65% thick fibres, 20%–35% thin fibres and small amounts of other types with the lumbosacral region having higher compositions of thick and thin fibres than those in the thoracodorsal region. Furthermore, the thoracodorsal and lumbosacral regions have the highest composition of thick fibre in the deeper dermis and quill follicle, respectively. These demonstrated that the complexity of the skin structure of Sunda porcupine due to its quill derivates correlated with its collagen composition and distribution.
... Old World porcupines have a wide geographical distribution between Africa and Eurasia with a broad latitudinal gradient (Mori et al., 2019). These herbivorous animals have quills covered in dense keratin plates, resembling spines, and are intricately integrated into the musculature of their skin (Chapman and Roze, 1997;Zherebtsova, 2000;Mori et al., 2014). During late Miocene, nine Hystrix species are recorded in Eurasia: H. parvae (Kretzoi, 1951), H. primigenia (Wagner, 1848) and H. depereti (S ßen, 2001a) in Europe; H. primigenia and H. kayae (Halaçlar et al., 2024) in Asia Minor; and H. lufengensis (Wang and Qi, 2005), H. gansuensis (Wang and Qiu, 2002), H. aryanensis (S ßen, 2001b), H. brevirostra (Wang and Qiu, 2020) and H. paukensis (Nishioka et al., 2011) in Asia (excluding Asia Minor) (Van Weers, 1990;Lopatin et al., 2003;Van Weers and Rook, 2003;S ßen and Purabrishemi, 2010;Flynn and Wu, 2017;Wang and Qiu, 2020;Halaçlar et al., in press). ...
This study introduces Hystrix aegeanensis n. sp., a new late Miocene (MN 12) porcupine species from Samos, Greece, and Karaağaç, Türkiye. This represents the fifth record of late Miocene Hystrix from Europe and Asia Minor. H. aegeanensis n. sp. exhibits a unique occlusal outline and pattern compared to those of all other late Miocene Hystrix species. This study further clarifies the systematic divergence among late Miocene Hystrix species based on occlusal outlines and patterns. The first group, including H. parvae, H. primigenia, H. gansuensis, H. brevirostra, and H. lufengensis, exhibits a simple occlusal pattern and an oval/square occlusal outline. In contrast, the second group, comprising H. aegeanensis n. sp., H. kayae, and H. aryanensis, displays a more complex occlusal pattern and a rectangular occlusal outline on their upper cheek teeth.
... Consequently, as already noted, the eigenvectors are the axes, and the eigenvalues index the lengths of these axes. Meanwhile, Gould 9 compares eigenfunctions for a large number of areal unit polygons to the quills covering a porcupine 10 , which number about 30,000 in adult animals and 157 range from one-half to four inches in length. This length variation (e.g., their eigenvalue analogy) arises in part because new quills grow to replace released ones, and in part because quills naturally grow to different lengths. ...
This exposition presents little-known connections between geography, through geographic information systems (GISs), mathematics, through matrix algebra, and art, through paintings and images, adding to the geo-humanities, spatial humanities, and humanistic mathematics literature. To this end, findings summarized for spatial statistical analyses of selected Susie Rosmarin paintings (which are reminiscent of visualizations of certain mathematical quantities known as eigenvectors), remotely sensed images that have appeared in art exhibits, and selected famous paintings by historically renowned artists reveal that spatial autocorrelation constitutes a fundamental element of art. These analyses extend the tradition of visualizing fractals as art, and interfacing cartography with art. This paper promotes analytical art, and establishes additional commonalities for GIScience, mathematics, and art.
... Quills and spines are structural modification of the hairs, which are found in some mammals, including porcupines, spiny mouse and rats, hedgehogs, tenrecs and echidnas (van Weers 1979(van Weers , 1983Zherebtsova 2000). Previous research on Hystrix javanica and Erethizon dorsatum, the new world porcupine, revealed that the sebaceous glands are also associated with the quill follicles, and have a larger size than those of the hair follicle (Prawira et al. 2018b;Chapman and Roze 1997). This suggests that the function of the glands to lubricate quills is greater than that of sebaceous glands in hair follicles. ...
Prawira AY, Novelina S, Farida WR, Darusman HS, Agungpriyono S. 2019. Lectin histochemical study of the quill sebaceous gland in the dorsal skin of the Sunda porcupine (Hystrix javanica). Biodiversitas 20: 2677-2684. In the Sunda Porcupine skin, the sebaceous glands of the quill follicles are multi-lobed alveolar and better developed than those of hair follicles. Using lectin histochemistry, we have studied the distribution of sugar-binding in the sebaceous glands of quills in comparison with those of hairs in six adult Sunda Porcupines. The skin samples from the thoracodorsal and lumbosacral regions were collected by biopsy procedure and processed for histology, histochemistry, and lectin histochemistry. The results showed that the lectin binding patterns are similar in secretory acinar parts of both types of sebaceous glands. The acini and excretory duct contained neutral carbohydrate and sugar residues. The acini also contained alpha D-mannose sugar residue, while the non-secretory excretory duct and debris in the lumen contained alpha-D-mannose, alpha-L-fucose, and α>β-N-acetylgalactosamine, and complex type N-glycan (oligosaccharide) sugar residues, as well. The present findings allowed us to suggest, that in Sunda Porcupine functions of the sebaceous glands of quills are more complex and active compared to those of the hairs.
... These fibers had a clear boundary with the dermal reticular layer in the dorsal thoracic region to the caudal radix and were visibly attached to the apex caudal region (Figure 2a and 2c). The density of woven fibers on the sheath of the thorn follicles in Sunda porcupine was similar to that of the follicle sheath structure of the Erethizon dorsatum (porcupine) consisting of a dense collagen bundle network in a circular arrangement (Chapman and Roze, 1997). The thickness of the connective tissue of the follicle of the thorns varies depending on the region and the type of thorn. ...
This study was conducted to investigate the histological characteristic, type, and distribution of connective tissue in Sunda porcupine skin. The investigation was carried out in three adult of sunda porcupines at microscopic level using hematoxylin eosin, Masson thrichrome, Verhoeffs van Gieson, alcian blue pH 2.5 and periodic acid Schiff staining methods. Skin consists of epidermis, dermis hypodermis, and subcutaneous muscle. Quill follicles were the main and dominant structure as well as the specific characteristic on Sunda porcupine skin. The connective tissue was distributed well in basal membrane, dermis, quill follicle, and hypodermis with various intensity and density. The collagen was the main fiber found in the skin while the elastin fiber was not observed. The acid carbohydrate was found distributed well in the skin while the neutral carbohydrate was not detected in this study. In addition the fibers of connective tissue associated with the adipose tissue which found plentifully in quill follicles and hypodermis. The present results showed that the wide distribution of connective tissue might have an important role on the wound healing physiology of Sunda porcupine skin.
Sarcoptic mange causes pruritic and crusting dermatitis in a large number of mammalian species with varying population impacts. Between 2016 and 2022, 15 North American porcupines (Erethizon dorsatum) were diagnosed with sarcoptic mange at Cornell University's Janet L. Swanson Wildlife Hospital in Ithaca, New York. Disease severity varied among individuals but all shared a similar unique presentation with thick, pale tan to yellow crusts limited in distribution to the ventral, nonquilled areas of the body, including the ventral abdomen and thorax, distal limbs, and face. The thick, hard nature of the crusts resulted in additional complications in many individuals, including inability to move the jaw and cracking and fissuring of the crusts and skin over joints of the limbs. Mites were plentiful within the crusts, with some burrowing into the epidermis as deep as the stratum spinosum. Secondary bacterial and/or fungal dermatitis were common, resulting in sepsis and death in three of the porcupines. Treatment with avermectins (ivermectin and/or selamectin) for 4–5 wk was successful in 12 cases in combination with other supportive care measures, including subcutaneous fluids, antimicrobials, and analgesics. Porcupines were hospitalized for an average of 18 d (ranging from 7 to 50 d) prior to transfer to a licensed wildlife rehabilitator for continued treatment and eventual release back into the wild.
Scanning electron microscopy shows that the quill surface from the thin-spined porcupine (Chaetomys subspinosus (Olfers, 1818)) has several regional cuticular patterns of which the most expansive is highly pitted with 4 μm wide pores leading to pits, which in turn communicate circumferentially via tunnels to neighboring pits. The cell unit of the pitted layer is a hexagonal or pentagonal prism, the “pitted fibrillous cuticular cell” (PFCC), which has a superficial pitted cuticular scale derived part with an underlying part packed with fine fibrils and the nucleus. The scaled part has about six scales, which swell and fuse except where raised cuticular scales from the inner root sheath intervene. This unique and complicated cell soon dies even while below the skin’s surface during its development. The raised scale-bearing cells of the inner root sheath leave their imprints on the PFCC’s surface, thus indicating that the quill’s pitted layer is at one time soft. Histochemistry, energy dispersive X-ray spectrometry, and Fourier transform infrared spectrometry show that the PFCC is keratinous.
Could a porcupine make a good pet? Do they ever stick themselves or other porcupines with their quills? In this latest addition to the Animal Answer Guide series, we learn about these mysterious animals' "pincushion defense" along with the following facts: porcupines survive on a diet of leaves, bark, and fruit quills are actually modified hairs there are 26 species of porcupines (and counting) Old World and New World porcupines have a common ancestor but evolved independently New World males will gather to fight ferociously over a single female Porcupines: The Animal Answer Guide presents solid, current science of porcupine biology. Uldis Roze compares porcupines in terms of body plan, behavior, ecology, reproduction, and evolutionary relationships. He examines the diversity of porcupines from around the world-from North and South America to Africa and Asia. This guide explores the interactions between humans and porcupines, including hunting, use of quills by aboriginal societies, efforts to poison porcupines, and human and pet injuries (and deaths) caused by porcupines. Roze also highlights the conservation issues that surround some porcupine species, such as the thin-spine porcupine of Brazil, which is so rare that it was thought to be extinct until its rediscovery in the 1980s. © 2012 The Johns Hopkins University Press. All rights reserved.
Phylogenetic analyses of cytochrome-b sequence data from 13 of the 15 currently recognized species of New World porcupines were used to test competing taxonomic hypotheses and to explore scenarios of morphological evolution and biogeography. Consistent with previous studies, the monophyly of Erethizontidae (Chaetomys + Erethizon + Coendou) and the monophyly of Erethizontinae (Erethizon + Coendou) were both strongly supported. However, cytochrome- b sequence data provide no support for the reciprocal monophyly of Coendou, “Echinoprocta,” and “Sphiggurus” as those taxa were previously recognized by authors. All of the erethizontid species recognized in recent revisionary work and represented by multiple sequences in this study were recovered as monophyletic groups. Maximum-likelihood (ML) analyses of these data recovered the following phylogeny for 11 species of Coendou: ((melanurus (ichillus (pruinosus + vestitus))) ((spinosus (bicolor + nycthemera)) (prehensilis (mexicanus (quichua + rufescens))))). Ancestral-state reconstructions based on the ML topology suggest that several morphological characters emphasized in past erethizontid classifications (size, nasofrontal sinus inflation, and long fur) have evolved homoplasiously Maximum-likelihood inference of geographic range evolution suggests that the last common ancestor of living erethizontids was a cis-Andean species, and that most subsequent cladogenesis was also cis-Andean; however, at least two trans-Andean dispersal events are plausibly indicated, as well as two separate invasions of Andean landscapes. Among the most remarkable results of this study are almost-identical sequences of Coendou prehensilis from localities spanning 27° of latitude and 25° of longitude; we speculate that a trophic-niche shift might have allowed rapid range expansion of this species, which accounts for almost all known cases of geographic range overlap and sympatry in the genus Coendou.
The interdigital glands of the moose (Alces alces andersoni) are distinguished from the neighboring skin by the lighter epidermis, green hairs, and huge sebaceous and sweat glands. In the glandular area is a shallow fore pocket and deep hind pocket in both sexes and all ages. Special 250 μm thick sections helped reveal the pilosebaceous–sweat gland unit in both the gland and general body surface. The secretory cycle of the apocrine cells and the cycle's relationship to the lipofuscin cycle are described. Macrophages ingest lipofuscin from moribund apocrine cells. The site of the dead apocrine cell is replaced by neighboring epithelial cells spreading out to fill the gap. The green hairs have a transmittance peak at 515 nm. The chemical nature of the coloring matter is unknown but is probably an organic compound covalently bonded to keratin. The green secretion of the sweat gland in some way dyes the hair. In both sexes the sebaceous gland volume of the gland increases at the rut.
The force required to epilate single chest hairs was measured electronically during a slow (0.09 m/s) machine-driven extraction. Telogen hairs showed a mean anchoring strength of 70 g (s.d. = 16). Anagen hairs, with variable amounts of surrounding root sheaths, had a mean of 71 g (s.d. = 10) whereas bare anagen hairs had a mean of 66 g (s.d. = 13). The catagen phase had a mean of 62 g (s.d. = 8). It is highly likely that the anchoring mechanisms of anagen and telogen roots are as different physiologically as these roots are different anatomically yet both are sufficiently strong to resist the ordinary tractions and pullings to which hair is subject.
Three objective parameters which characterize the main features of the epidermis are described. One parameter relates to the mean thickness of the epidermis determined by a simple method. The second descriptive number is the ratio of the thickness of the strata corneum, lucidum and granulosum to the stratum matpighii. The third number relates to the degree of folding of the dermo-epidermal junction.
The ratio of the mean thickness of abnormal to normal stratum corneum (for the same skin region) is suggested as an objective measure of hyperkeratosis; likewise, the ratio of abnormal to normal stratum malpighii is an objective measure of acanthosis.
A single parameter can be measured against time to follow changes during the pathogenesis, treatment or remission of epidermal disease.
As psoriasis undergoes progressive acanthosis, the geometry of the folding of the dermo- epidermal junction is such that cells of the stratum malpighii retain their close association with the cylindrical dermal papillae.