Anal gland

Last updated

The anal glands or anal sacs are small glands near the anus in many mammals. [1] They are situated in between the external anal sphincter muscle and internal anal sphincter muscle. [2] Their function in humans is unclear. [3]

Contents

Sebaceous glands within the lining secrete a liquid that is used for identification of members within a species. These sacs are found in many species of carnivorans, [2] including wolves, [4] bears, [5] [6] sea otters [7] and kinkajous. [8]

Anatomy

Diagram showing anal canal, with dentate line, along which anal crypts open. Anal glands drain into anal crypts via anal ducts. Note also intersphincteric plane where some of these glands are located. Gray1080.png
Diagram showing anal canal, with dentate line, along which anal crypts open. Anal glands drain into anal crypts via anal ducts. Note also intersphincteric plane where some of these glands are located.

The anal glands are situated within the wall of the anal canal [9] and communicate with the lumen of the canal via ducts that open at the anal valves, just proximal to the pectinate line. Humans have 12 anal glands on average (which are evenly distributed around the circumference of the anal canal). [3] The glands are situated at varying depths in the wall of the anal canal; some between the layers of the internal and external sphincter (the intersphincteric plane). [10] [11]

Function

In humans, the glands secrete mucin (which differs in composition from that secreted by the rectal mucosa). Their function is unclear. [3]

Clinical significance

Anal glands are the most common cause of anal fistula. Large fistulae present a surgical challenge as resection of larger sections of the anal sphincter may result in anal incontinence. [3] The cryptoglandular theory states that obstruction of these ducts[ clarification needed ], presumably by accumulation of foreign material (e.g. fecal bacterial plugging) in the crypts, may lead to perianal abscess and fistula formation. [10] [11]

Entry of bacteria into the lumen of the glands can cause infection (which may then spread), and inflammation can prevent drainage of the glands. [12]

Other animals

Dogs and cats

Anal gland abscess on a dog Anal gland abscess.JPG
Anal gland abscess on a dog

Dogs and cats primarily use their anal gland secretions to mark their territory, and generally will secrete small amounts of fluid every time they defecate. Many will often express these glands when anxious or frightened as well. Dogs who are healthy can usually have a wide variety in the appearance of their sac's content. [13] Anal sac fluid varies from yellow to tan or brown in color. The consistency of the fluid ranges from thin, watery secretions to thick, gritty paste. There can also be a range in the malodorousness, or how strongly the contents smell. [13] These factors can all be different from animal to animal. [13]

The inability to effectively express this fluid can lead to anal sacculitis. This is characterized by a build-up of fluid in the anal sac, an uncomfortable condition that can lead to pain and itching. Dogs and cats of any age may be affected, but dogs are far more likely to suffer from anal sacculitis than cats. Dogs and cats with anal glands that do not express naturally may exhibit specific signs, such as scooting the backside upon the ground, straining to defecate, and excessive licking of the anus. Cats may also defecate in areas outside the litter box. [14]

Discomfort may also be evident with impaction or infection of the anal glands. Anal sac impaction results from blockage of the duct leading from the gland to the opening. The sac is usually non-painful and swollen. Anal sac infection results in pain, swelling, and sometimes abscessation and fever.

Initial treatment usually involves the manual expression of the anal sacs, most often by a veterinary professional. The frequency of this procedure depends on the patient's individual degree of discomfort but can range from weekly to every few months. [14] Treatment may include lancing of an abscess or antibiotic infusion into the gland in the case of infection. The most common bacterial isolates from anal gland infection are E. coli , Enterococcus faecalis , Clostridium perfringens , and Proteus species. [15] Increasing dietary fiber is also commonly recommended. [14]

Anal sacs may be removed surgically in a procedure known as anal sacculectomy. This is usually done in the case of recurrent infection or because of the presence of an anal sac adenocarcinoma, a malignant tumor. Potential complications include fecal incontinence (especially when both glands are removed), tenesmus from stricture or scar formation, and persistent draining fistulae. [16]

Opossums

Opossums use their anal glands when they "play possum". As the opossum mimics death, the glands secrete a foul-smelling liquid, suggesting the opossum is rotting. Opossums are not members of the carnivora, and their anal sacs differ from those of dogs and their relatives. [8]

Skunks

Skunks use their anal glands to spray a foul-smelling and sticky fluid as a defense against predators.[ citation needed ]

Symbiotic relationships

Beavers

Both female and male beavers (Castor canadensis) have a pair of castor sacs and a pair of anal glands between the pelvis and tail. The physiology of the castor sac is unique to the beaver and contains an outer, vascular layer of connective tissue, a thicker layer of epithelial cells, and an inner packed layer of epithelial sheets. [17] Castoreum is the chemical compound that is secreted from the castor sacs and is originally a thin, yellowish liquid. It is composed of a mixture of varied metabolic compounds, from the sacs and other bodily systems, and excreted urine. [17] An extraction of castoreum, the scent glands from the male and female beaver are used in perfumery and as a flavor ingredient.

The vestibule of the anal glands are connected to the ducts of the castor glands. The anal glands of beavers are characterized as holocrine sebaceous glands, which means they secrete substances by disintegrating cells in the process. [18] Variations in color of anal glands range from a light straw color to brown. Compared to the castor sacs, anal glands secrete a much more sharp odor. [18] Beavers do have a presence of bacterial flora in their anal glands, with the most abundant being B. fragilis and E. coli. These two bacteria are common in the digestive tracts of mammals and are seen across all beaver populations regardless of sex, colony, age, class, and other factors. [18] This gives evidence to the idea that beavers do not have varied bacterial flora or significant varied odors within a family. Furthermore, this opposes the bacterial fermentation hypothesis that is common among a number of mammalian species. [17]

Beavers create scent mounds, which are essentially “mud pies”, where they deposit castoreum and other secretions on top of them. As scent communication is a common method across many species, beavers use scent mounds as a way to alert that a region is inhabited by a family of beavers and to mark territory. [19] The frequency of scent mounding is typically highest during the spring and after winter when the ice has melted. This is prevalent as the water sources are more available from which beavers can gather mud. [19] Scent mounding is highest during intergroup interactions, vicinity of abutting beaver populations, and gestation periods. [19] Scent mounds are mainly a medium of communication against adjacent beaver populations and it prevents the exploitation of food resources, marks distinct family territories, and prevents extreme colonization of a habitat. [19]

Badgers

The European badger (Meles meles), a nocturnal carnivore, has a powerful olfactory system. [20] Anal gland secretions (AGS) of badgers are not commonly studied but contribute to key information for communication due to volatile organic compounds (VOCs). [20] Like many other carnivorous mammals, such as mongoose, bears, and otters, they can detect sex differences from the different ratios of compounds of VOCs. [20] VOCs in AGS of badger samples have individual-specific information, including health, fitness, reproductive status, and group membership. [20] Male badgers react differently to VOCs in AGS from fertile females versus non-fertile females. [20] The VOCs can change yearly based on diet and environmental factors, and seasonal changes of VOCs are influenced by breeding season. [20]

The subcaudal gland is right next to the anal sac. [21] The subcaudal gland contributes to individual-specific communication, much like the AGS. [21] Subcaudal glands of badgers had 56 operational taxonomic units (OTU) used to classify the different bacteria found into four different phyla. [21] Based on a study by Yung Wa Sin 2012, conducted on 79 subcaudal secretions from summer and spring, OTUs fall into four bacterial phyla: Actinobacteria, Firmicutes, Proteobacteria, and Bacteroidetes. [21] Actinobacteria was the dominant phyla as it represented >76% of all bacterial communities in the badger adults. [21] Cub secretion microbial communities were significantly more diverse; Firmicutes were the most abundant bacterial phyla in adult badger microbial communities. [21] This bacterial dominance shift may be due to puberty in the cubs. [21] In the spring, a breeding female versus a non-breeding female had significant microbial community differences, but they did not see significant differences in bacterial communities in the summer months. [21] Secretions from the subcaudal gland are shown to be rich in short and medium-chain fatty acids likely produced by pheromone active products from actinobacteria long-chain fatty acids. [21]

In the majority of mammals, group integration is performed by the adult. However, in badgers cubs begin this gradual process (14–16 weeks). [22] Badgers have the reputation of being aggressive towards one another and cubs are often victims to infanticide. [22] To lower aggression within a group of badgers, members will allogroom (a form of social grooming) or allomark (transferring scents between other group members) more frequent and intense during spring months. [22] Maturing cubs rub themselves against adult badger's subcaudal region via allomarking. [22] This is also referred to as “scent theft” as cubs that do this have the same group scent. [22] Badger cubs do not have the subcaudal gland secretion ability until approximately four months old . [22]

Wolves

Volatile compounds found in the anal sac secretions of intact males, intact females, castrate males, ovariectomized females, and anosmic/ pinealectomized males and females were analyzed using gas chromatography. [20] The volatile compounds found in the anal-gland secretions were largely alcohols, aldehydes, and ketones. [20] Of the volatile compounds that were analyzed, the relative quantities of volatile compounds present in the test subjects’ anal sac secretions varied significantly between secretions collected outside and during mating season. [20] This indicates that the volatile compounds in the anal sac secretions are used to signal information like gender and reproductive status. [20] Results of this study indicated that some of the volatile compounds, specifically 2-octenal, 2-octen-1-ol and indole, were produced by microbes. [20]

Indian mongoose

The anal gland of the Indian mongoose consists of large sebaceous glands that surround the anus, called the anal sac. [23] The anal sac remains covered in sebum and remains closed when the tail of the mongoose is down but opens when the tail is raised. [23] Inside the anal sacs are the ducts of the two anal pockets that lie on either side of the anus. [23] The mongoose marks objects in its habitat by rubbing that object with the anal area leaving behind the distinct scent of carboxylic acids. [23] Contents of the anal pocket secretions revealed 6 saturated carboxylic acids: acetic, propionic, isobutyric, butyric, isovaleric and valeric. [23] There are currently no notable differences in the chemicals found in the anal pocket secretions between the sexes. [23]

Evidence shows that the carboxylic acids found in the anal pocket secretions are produced by bacterial metabolism of contents inside the sebum. [23] The bacteria isolated from anal sac secretions have been identified as Peptococcusspp., Peptostreptococcus plagarumbelli, Bacillus cereus and Eubacterium or Catenobacteriumspp. [23] These bacterial species have been found to produce carboxylic acids within the anal pocket. [23]

Hyenas

Hyenas are known to engage in “pasting” which is a type of scent marking behavior. This paste is rich in lipid sebum and epithelial cells, and is produced by sebaceous glands which then go directly into the anal glands and on top of a grass stalk. [24] The organ that provides this paste, the anal glands, are occupied by microbes. Although both species of hyena contain fermentative bacteria, the microbes found in the anal gland of spotted hyenas (Crocuta crocuta) differ from the microbes found in striped hyenas. [25]

The spotted hyena paste holds many populations of coccus and rod shaped bacteria. A survey found that Firmicutes, Actinobacteria, Bacteroidetes, and Proteobacteria are the most common bacteria found in the hyena paste. [24] Although many bacteria have been identified, around half are still unidentified. [24]

Microbiome populations also vary by sex and age. The adult and juvenile hyenas’ anal glands have the least diverse microbiota of their whole body. For females, the most common bacteria found in their paste was Anaerococcus, Anaerovorax, Corynebacterium, Eubacterium, Helcococcus, Porphyromonas, and Propionibacterium. [24] Compared to the male hyenas which have a different microbiota than female hyenas. Juvenile males have more Prevotella and Firmicutes, while juvenile females have more Corynebacterium and Clostridiales. There is also a difference between adult female hyenas and juvenile hyenas. Juvenile female hyenas have more Erysipelotrichaceae and Helicobacter than the adult hyenas. These bacteria are common to the milk that hyenas feed their young. [26]

Dogs and domestic cats

The bacteria found in the feces and anal glands of dogs and cats are also found in their mouths due to the consistent exposure to their backsides (licking and chewing). [27] In a healthy dog or cat the bacteria normally found in their feces are Streptococci and Enterococci, more specifically, Enterococcus faecium, Streptococcus bovis, and Enterococcus faecalis. [27]

Dogs have two anal sacs which are located in the connective tissue off the anus. [13] There are many glands in dogs that secrete into the sac's lumen, filling them with fluid. [13] The anal-sacs are usually made up of about 88% water, 11.5% organic and 0.5% inorganic matter. [13] The secretion of anal gland content contains mucin, that is rich in sialic acid and other anti-microbial proteins, like lysozyme, immunoglobulin A, and lactoferrin. [13] There is also an abundance of gram-positive cocci since they are part of the anal glands normal flora. [13] The organic components mainly consist of short-chain fatty acids and trimethylamine. [13] The anal-sac fluid can be secreted or expressed into the anal channel to serve as a scent marker for an individual's territory through their feces. [13]

See also

Related Research Articles

<span class="mw-page-title-main">Beaver</span> Genus of semiaquatic rodents that build dams and lodges

Beavers are large, semiaquatic rodents of the Northern Hemisphere. There are two existing species: the North American beaver and the Eurasian beaver. Beavers are the second-largest living rodents, after capybaras, weighing up to 50 kg (110 lb). They have stout bodies with large heads, long chisel-like incisors, brown or gray fur, hand-like front feet, webbed back feet, and tails that are flat and scaly. The two species differ in skull and tail shape and fur color. Beavers can be found in a number of freshwater habitats, such as rivers, streams, lakes and ponds. They are herbivorous, consuming tree bark, aquatic plants, grasses and sedges.

<span class="mw-page-title-main">Skunk</span> Common name of mammals in the family Mephitidae

Skunks are mammals in the family Mephitidae. They are known for their ability to spray a liquid with a strong, unpleasant scent from their anal glands. Different species of skunk vary in appearance from black-and-white to brown, cream or ginger colored, but all have warning coloration.

Digestion is the breakdown of large insoluble food compounds into small water-soluble components so that they can be absorbed into the blood plasma. In certain organisms, these smaller substances are absorbed through the small intestine into the blood stream. Digestion is a form of catabolism that is often divided into two processes based on how food is broken down: mechanical and chemical digestion. The term mechanical digestion refers to the physical breakdown of large pieces of food into smaller pieces which can subsequently be accessed by digestive enzymes. Mechanical digestion takes place in the mouth through mastication and in the small intestine through segmentation contractions. In chemical digestion, enzymes break down food into the small compounds that the body can use.

<span class="mw-page-title-main">Saliva</span> Bodily fluid secreted by salivary glands

Saliva is an extracellular fluid produced and secreted by salivary glands in the mouth. In humans, saliva is around 99% water, plus electrolytes, mucus, white blood cells, epithelial cells, enzymes, antimicrobial agents.

<span class="mw-page-title-main">Sebaceous gland</span> Gland to lubricate the hair and skin

A sebaceous gland or oil gland is a microscopic exocrine gland in the skin that opens into a hair follicle to secrete an oily or waxy matter, called sebum, which lubricates the hair and skin of mammals. In humans, sebaceous glands occur in the greatest number on the face and scalp, but also on all parts of the skin except the palms of the hands and soles of the feet. In the eyelids, meibomian glands, also called tarsal glands, are a type of sebaceous gland that secrete a special type of sebum into tears. Surrounding the female nipple, areolar glands are specialized sebaceous glands for lubricating the nipple. Fordyce spots are benign, visible, sebaceous glands found usually on the lips, gums and inner cheeks, and genitals.

<span class="mw-page-title-main">Spotted hyena</span> Species of hyena

The spotted hyena, also known as the laughing hyena, is a hyena species, currently classed as the sole extant member of the genus Crocuta, native to sub-Saharan Africa. It is listed as being of least concern by the IUCN due to its widespread range and large numbers estimated between 27,000 and 47,000 individuals. The species is, however, experiencing declines outside of protected areas due to habitat loss and poaching. Populations of Crocuta, usually considered a subspecies of Crocuta crocuta, known as cave hyenas, roamed across Eurasia for at least one million years until the end of the Late Pleistocene. The spotted hyena is the largest extant member of the Hyaenidae, and is further physically distinguished from other species by its vaguely bear-like build, rounded ears, less prominent mane, spotted pelt, more dual-purposed dentition, fewer nipples, and the presence of a pseudo-penis in the female. It is the only placental mammalian species where females lack an external vaginal opening, having a pseudo-penis instead.

<span class="mw-page-title-main">Areolar gland</span> Oil glands around the nipple that promote breastfeeding

Areolar glands, also known as glandulae areolares, Montgomery glands, and tubercula areolae, are 10-15 elevations found on the areola. They are usually arranged in a circle around the nipple, and can be particularly visible when the nipple is erect. Their role is to promote adequate breastfeeding of the infant.

Body odor or body odour (BO) is present in all animals and its intensity can be influenced by many factors. Body odor has a strong genetic basis, but can also be strongly influenced by various factors, such as sex, diet, health, and medication. The body odor of human males plays an important role in human sexual attraction, as a powerful indicator of MHC/HLA heterozygosity. Significant evidence suggests that women are attracted to men whose body odor is different from theirs, indicating that they have immune genes that are different from their own, which may produce healthier offspring.

<span class="mw-page-title-main">Sebacic acid</span> Chemical compound

Sebacic acid is a naturally occurring dicarboxylic acid with the chemical formula HO2C(CH2)8CO2H. It is a white flake or powdered solid. Sebaceus is Latin for tallow candle, sebum is Latin for tallow, and refers to its use in the manufacture of candles. Sebacic acid is a derivative of castor oil.

<span class="mw-page-title-main">Sweat gland</span> Small sweat-producing tubular skin structures

Sweat glands, also known as sudoriferous or sudoriparous glands, from Latin sudor 'sweat', are small tubular structures of the skin that produce sweat. Sweat glands are a type of exocrine gland, which are glands that produce and secrete substances onto an epithelial surface by way of a duct. There are two main types of sweat glands that differ in their structure, function, secretory product, mechanism of excretion, anatomic distribution, and distribution across species:

<span class="mw-page-title-main">Castoreum</span> Fluid produced by beavers

Castoreum is a yellowish exudate from the castor sacs of mature beavers. Beavers use castoreum in combination with urine to scent mark their territory. Both beaver sexes have a pair of castor sacs and a pair of anal glands, located in two cavities under the skin between the pelvis and the base of the tail. The castor sacs are not true glands on a cellular level, hence references to these structures as preputial glands, castor glands, or scent glands are misnomers.

<span class="mw-page-title-main">Scent gland</span> Exocrine glands found in most mammals

Scent gland are exocrine glands found in most mammals. They produce semi-viscous secretions which contain pheromones and other semiochemical compounds. These odor-messengers indicate information such as status, territorial marking, mood, and sexual behaviour. The odor may be subliminal—not consciously detectable. Though it is not their primary function, the salivary glands may also function as scent glands in some animals.

<span class="mw-page-title-main">Violet gland</span> Gland on the tail of certain mammals

The violet gland or supracaudal gland is a gland located on the upper surface of the tail of certain mammals, including European badgers and canids such as foxes, wolves, and the domestic dog, as well as the domestic cat. Like many other mammalian secretion glands, the violet gland consists of modified sweat glands and sebaceous glands.

Dogs, as with all mammals, have natural odors. Natural dog odor can be unpleasant to dog owners, especially when dogs are kept inside the home, as some people are not used to being exposed to the natural odor of a non-human species living in proximity to them. Dogs may also develop unnatural odors as a result of skin disease or other disorders or may become contaminated with odors from other sources in their environment.

An apocrine sweat gland is composed of a coiled secretory portion located at the junction of the dermis and subcutaneous fat, from which a straight portion inserts and secretes into the infundibular portion of the hair follicle. In humans, apocrine sweat glands are found only in certain locations of the body: the axillae (armpits), areola and nipples of the breast, ear canal, eyelids, wings of the nostril, perineal region, and some parts of the external genitalia. Modified apocrine glands include the ciliary glands in the eyelids; the ceruminous glands, which produce ear wax; and the mammary glands, which produce milk. They are distinct from eccrine sweat glands which cover the whole body.

Gastrointestinal physiology is the branch of human physiology that addresses the physical function of the gastrointestinal (GI) tract. The function of the GI tract is to process ingested food by mechanical and chemical means, extract nutrients and excrete waste products. The GI tract is composed of the alimentary canal, that runs from the mouth to the anus, as well as the associated glands, chemicals, hormones, and enzymes that assist in digestion. The major processes that occur in the GI tract are: motility, secretion, regulation, digestion and circulation. The proper function and coordination of these processes are vital for maintaining good health by providing for the effective digestion and uptake of nutrients.

Margaric acid, or heptadecanoic acid, is a saturated fatty acid. Its molecular formula is CH3(CH2)15CO2H. Classified as an odd-chain fatty acid, it occurs as a trace component of the fat and milkfat of ruminants. Salts and esters of margaric acid are called heptadecanoates.

<span class="mw-page-title-main">Male accessory gland</span> Sexual gland in males

Male accessory glands (MAG) in humans are the seminal vesicles, prostate gland, and the bulbourethral glands. In insects, male accessory glands produce products that mix with the sperm to protect and preserve them, including seminal fluid proteins. Some insecticides can induce an increase in the protein content of the male accessory glands of certain types of insects. This has the unintended effect of increasing the number of offspring they produce.

<span class="mw-page-title-main">Human digestive system</span> Digestive system in humans

The human digestive system consists of the gastrointestinal tract plus the accessory organs of digestion. Digestion involves the breakdown of food into smaller and smaller components, until they can be absorbed and assimilated into the body. The process of digestion has three stages: the cephalic phase, the gastric phase, and the intestinal phase.

The biochemistry of body odor pertains to the chemical compounds in the body responsible for body odor and their kinetics.

References

  1. Howard E. Evans; Alexander de Lahunta (7 August 2013). Miller's Anatomy of the Dog - E-Book. Elsevier Health Sciences. ISBN   978-0-323-26623-9.
  2. 1 2 Estes R (1992). The Behavior Guide to African Mammals: Including Hoofed Mammals, Carnivores, Primates. University of California Press. ISBN   978-0-520-08085-0.
  3. 1 2 3 4 Standring, Susan (2020). Gray's Anatomy: The Anatomical Basis of Clinical Practice (42th ed.). New York. p. 1201. ISBN   978-0-7020-7707-4. OCLC   1201341621.{{cite book}}: CS1 maint: location missing publisher (link)
  4. L. David Mech; Luigi Boitani (1 October 2010). Wolves: Behavior, Ecology, and Conservation. University of Chicago Press. ISBN   978-0-226-51698-1.
  5. Rosell F, Jojola SM, Ingdal K, Lassen BA, Swenson JE, Arnemo JM, Zedrosser A (Feb 2011). "Brown bears possess anal sacs and secretions may code for sex" (PDF). Journal of Zoology. 283 (2): 143–152. doi:10.1111/j.1469-7998.2010.00754.x. hdl: 11250/2437930 .
  6. Dyce KM, Sack WO, Wensing CJ (1987). Textbook of Veterinary Anatomy. W.B. Saunders Company. ISBN   0-7216-1332-2.
  7. Kenyon KW (1969). The Sea Otter in the Eastern Pacific Ocean. Washington, D.C.: U.S. Bureau of Sport Fisheries and Wildlife.
  8. 1 2 Ford LS, Hoffman RS (1988-12-27). "Potos flavus". Mammalian Species (321). American Society of Mammalogists: 1–9. doi: 10.2307/3504086 . JSTOR   3504086.
  9. Pfenninger, John L.; Zainea, George G. (15 June 2001). "Common Anorectal Conditions: Part I. Symptoms and Complaints". American Family Physician. 63 (12): 2391–2398. PMID   11430454.
  10. 1 2 Yamada T, Alpers DH, Kalloo AN, Kaplowitz N, Owyang C, Powell DW, eds. (2009). Textbook of gastroenterology (5th ed.). Chichester, West Sussex: Blackwell Pub. ISBN   978-1-4051-6911-0.
  11. 1 2 Wolff BG, Pemberton JH, Wexner SD, Fleshman JW, Beck DE, eds. (2007). The ASCRS textbook of colon and rectal surgery. New York: Springer. ISBN   978-0-387-24846-2.
  12. Standring, Susan (1201). Gray's Anatomy: The Anatomical Basis of Clinical Practice (42th ed.). New York. p. 683. ISBN   978-0-7020-7707-4. OCLC   1201341621.
  13. 1 2 3 4 5 6 7 8 9 10 Beynen AC (1985-03-01). "Biochemists and the diet of their rats". Trends in Biochemical Sciences. 10 (3): 108–109. doi:10.1016/0968-0004(85)90265-8. ISSN   0968-0004.
  14. 1 2 3 "Anal sacculitis in dogs | Vetlexicon Canis from Vetstream | Definitive Veterinary Intelligence". www.vetstream.com. Retrieved 2019-12-05.
  15. Ettinger SJ, Feldman EC (1995). Textbook of Veterinary Internal Medicine (4th ed.). W.B. Saunders Company. ISBN   0-7216-6795-3.
  16. Hill LN, Smeak DD (September 2002). "Open versus closed bilateral anal sacculectomy for treatment of non-neoplastic anal sac disease in dogs: 95 cases (1969-1994)". Journal of the American Veterinary Medical Association. 221 (5): 662–665. doi: 10.2460/javma.2002.221.662 . PMID   12216905.
  17. 1 2 3 Walro JM, Svendsen GE (May 1982). "Castor sacs and anal glands of the north american beaver (Castor canadensis): their histology, development, and relationship to scent communication". Journal of Chemical Ecology. 8 (5): 809–819. doi:10.1007/BF00994781. PMID   24415179. S2CID   19327243.
  18. 1 2 3 Svendsen GE, Jollick JD (1978). "Bacterial contents of the anal and castor glands of beaver (Castor canadensis)". Journal of Chemical Ecology. 4 (5): 563–569. doi:10.1007/BF00988920. ISSN   0098-0331. S2CID   25447438.
  19. 1 2 3 4 Svendsen GE (1980). "Patterns of scent-mounding in a population of beaver (Castor canadensis)". Journal of Chemical Ecology. 6 (1): 133–148. doi:10.1007/BF00987533. ISSN   0098-0331. S2CID   22234085.
  20. 1 2 3 4 5 6 7 8 9 10 11 Raymer J, Wiesler D, Novotny M, Asa C, Seal US, Mech LD (May 1985). "Chemical investigations of wolf (Canis lupus) anal-sac secretion in relation to breeding season". Journal of Chemical Ecology. 11 (5): 593–608. doi:10.1007/BF00988570. PMID   24310125. S2CID   26151726.
  21. 1 2 3 4 5 6 7 8 9 Gorman M, Nedwell DB, Smith RM (1974). "An analysis of the contents of the anal scent pockets of Herpestes auropunctatus (Carnivora: Viverridae)". Journal of Zoology. 172 (3): 389–399. doi:10.1111/j.1469-7998.1974.tb04115.x. ISSN   0952-8369.
  22. 1 2 3 4 5 6 Sin YW, Buesching CD, Burke T, Macdonald DW (September 2012). "Molecular characterization of the microbial communities in the subcaudal gland secretion of the European badger (Meles meles)". FEMS Microbiology Ecology. 81 (3): 648–659. doi: 10.1111/j.1574-6941.2012.01396.x . PMID   22530962.
  23. 1 2 3 4 5 6 7 8 9 Gorman M, Nedwell DB, Smith RM (1974). "An analysis of the contents of the anal scent pockets of Herpestes auropunctatus (Carnivora: Viverridae)". Journal of Zoology. 172 (3): 389–399. doi:10.1111/j.1469-7998.1974.tb04115.x. ISSN   0952-8369.
  24. 1 2 3 4 Theis KR, Schmidt TM, Holekamp KE (2012-08-30). "Evidence for a bacterial mechanism for group-specific social odors among hyenas". Scientific Reports. 2 (1): 615. Bibcode:2012NatSR...2E.615T. doi:10.1038/srep00615. PMC   3431069 . PMID   22937224.
  25. Theis KR, Venkataraman A, Dycus JA, Koonter KD, Schmitt-Matzen EN, Wagner AP, et al. (December 2013). "Symbiotic bacteria appear to mediate hyena social odors". Proceedings of the National Academy of Sciences of the United States of America. 110 (49): 19832–19837. Bibcode:2013PNAS..11019832T. doi: 10.1073/pnas.1306477110 . PMC   3856825 . PMID   24218592.
  26. "Body site-specific microbiota reflect sex and age-class among wild spotted hyenas" . Retrieved 2022-04-25.
  27. 1 2 Devriese LA, Cruz Colque JI, De Herdt P, Haesebrouck F (November 1992). "Identification and composition of the tonsillar and anal enterococcal and streptococcal flora of dogs and cats". The Journal of Applied Bacteriology. 73 (5): 421–425. doi:10.1111/j.1365-2672.1992.tb04998.x. PMID   1447058.