Information for Vets GASTROINTESTINAL PHYSIOLOGY AND DISEASE IN THE DOMESTIC PET RABBIT by Susan A. Brown, DVM An inappropriate diet may be the most common factor linked to disease in the domestic pet rabbit. Dietary deficiencies in the rabbit can contribute to a wide range of conditions, including dental, urogenital, cardiac and infectious diseases and behavioural disorders. The direct effects of dietary deficiencies on the gastrointestinal tract (GIT) are an extremely common cause of enteritis and obstructive GIT disease. Therefore it is important to understand basic GIT physiology of this species, match this information to the lifestyle of the pet rabbit, and formulate a healthy diet. There are many resources available that discuss the GIT physiology of the domestic rabbit in great detail. An overview of this subject is presented below, and the reader is encouraged to look further for more detailed information. GASTROINTESTINAL PHYSIOLOGY Rabbits are strict herbivores, and their relatively small body size makes it difficult for them to store large volumes of coarse fibre. The indigestible fibre (in particular, lignocellulose) in the diet is eliminated from the GIT quickly, which thus controls the rate of peristalsis. The digestible portion of the diet is utilised to produce nutrients, some of which are absorbed directly through the wall of the GIT and some of which are reingested, digested and absorbed in the form of caecotropes. This particular system allows for a large volume of food intake with a rapid digestive transit time, increasing the total amount of energy stored and minimising the need to store fibre. The reingestion of nutrient-rich caecotropes is an important adaptation developed by wild lagomorph ancestors to allow survival in the face of an unreliable food supply.1 Oral Cavity The oral cavity, which marks the beginning of the GIT, contains open-rooted incisors and molars. Proper wear of the teeth depends a great deal on the type of diet that is fed. Wild rabbits take in a high proportion of tough, fibrous foods, which require a great deal of chewing. Diets that require little chewing result in inadequate tooth wear. 2 An example of this is commercial pellet diets, which are composed of tightly compressed small particles. Pellets can be pulverised into a size that can be swallowed with a minimum of chewing. Feeding a diet that is lacking in sufficient "chewing material" can lead to elongation of the crowns and/or the roots of the teeth.2 Overgrown crowns can result in enamel spurs on the molars and malocclusion of the incisors and/or molars. Hyperextended roots can lead to epiphora (from incisor roots compressing the lacrimal duct) and abscesses (as the roots penetrate the mandible or maxilla).2 Dental disease is a common cause of anorexia in the rabbit due to the pain associated with these conditions. In rabbits with suspected dental disease it is important to examine the mouth thoroughly and take high detail skull radiographs. Dental disease may be one of the most underdiagnosed causes of anorexia in the pet rabbit. Stomach Ingesta next enter the simple stomach with its small, nondistensible lumen and muscular pyloric sphincter. Rabbits are unable to vomit. The pH of the stomach is approximately 1 to 2 in the healthy adult rabbit.3 The pH of the rabbit stomach preweaning is closer to neutral (5 to 6.5), and the gut is lacking in bacterial flora until about 4 to 6 weeks of age.4 After this time the gut gradually becomes populated with living microorganisms and the pH of the stomach drops to 1 to 2. The exact gastric pH in all states of disease is currently unknown but appears to increase in the face of GIT disorders.5 A stomach food ball that contains sufficient fibre will have an "open network" composition and can be easily penetrated by the gastric fluids, which then sterilise the contents. A food ball that is composed of small particles that are compacted tightly, such as with a diet high in alfalfa meal or commercial pellets, may not be well penetrated and may allow passage of potential pathogens into the GIT.6 In addition, feeding sufficient fibre promotes healthy peristalsis, aiding the rapid movement of material out of the stomach and reducing the introduction of pathogens. A healthy rabbit’s stomach always contains ingesta mixed with hair (due to normal grooming). It is not possible to remove the food ball entirely by fasting. Instead, prolonged fasting may lead to the dehydration and impaction of the food ball as well as hepatic lipidosis.7 Gastric Stasis ("Trichobezoars") The condition commonly referred to as "gastric trichobezoars" or "hairballs" is a misnomer in my opinion and that of others.7 The problem is primarily related to a chronic gastric hypomotility resulting over time in an impacted, dehydrated food ball that cannot pass out of the stomach. Since hair is always present in the rabbit stomach, when the food ball becomes dehydrated, the hair becomes grossly more prominent, leading to the conclusion that the original problem was hair. The onset of gastric stasis is usually gradual, with a decrease in appetite and output of faeces; once complete stasis occurs, however, the rabbit becomes anorectic. Anorectic patients may still be active and alert for several days before the condition develops into a more severe and life-threatening GIT ileus. The most common cause of chronic gastric hypomotility is low dietary fibre, but one or more other factors can be involved including other dietary imbalances, inadequate water intake, inactivity, obesity, and concurrent disease. The classic sign of gastric stasis is a full stomach with a halo of air around very dense ingesta. In the past gastrotomy was performed in these cases, which often resulted in a high fatality rate. The current recommended treatment for gastric stasis is aggressive medical therapy that includes rehydrating the patient as well as the food ball, introducing high fibre force-feedings (and ultimately correcting the diet long term), giving intestinal motility stimulants, and providing other supportive care. The majority of mild to moderately affected cases respond to medical therapy within 72 hours. Cases that do not respond in this time need to be reevaluated as possible surgical candidates, but their prognosis is guarded to poor. Laxatives and proteindigesting enzymes such as pineapple juice have been recommended, but the use of these products alone does not correct the underlying problem and I do not find them useful. There is concern that long-haired rabbit breeds may accumulate an excessive amount of hair in the stomach during seasonal moults. In my experience these breeds do not exhibit gastric obstruction or stasis with any more frequency than any other breed if they are maintained yearround on a high fibre diet. Caecum and Colon Ingesta pass from the stomach into the small intestine, where they move rapidly toward the ileocaecocolonic junction. At this point the ingesta enter the hindgut and move into the caecum and the proximal colon. Through the contractions of muscular sacculations (haustra) in the walls of the caecum and proximal colon, the indigestible fibre and digestible material are efficiently sorted.3 Indigestible fibre migrates toward the centre of the intestinal lumen and is moved distally, forming the hard faeces. Digestible material settles into the haustra and is moved into the ceacum by reverse peristalsis of the proximal colon aided by fluid secretion into the colon.3 The large, well-developed caecum is a blind sac containing anaerobic gram-negative bacteria, including a predominant population of the bacilli Bacteroides.3 Many other organisms have been isolated, including potentially pathogenic bacteria such as Clostridium spiroforme and Escherichia coli, various protozoa, and a rabbit-specific yeast in the Saccharomycetaceae family.8 These organisms are not pathogenic if their populations remain small; however, any disruption of caecal environment (pH, motility, fluid content, dietary starch content, etc.) can result in a floral imbalance, leading to enteric disorders. Lactobacillus spp. are not commonly isolated in the rabbit caecum; therefore using oral Lactobacillus products in rabbits is of questionable therapeutic value. In addition, Lactobacillus does not adhere to or colonise the intestinal wall and is likely killed in the acid pH of the stomach or by the bile before it even reaches the hindgut.7 Through fermentation of the digestible material, caecal flora produce amino acids, fatty acids, and certain vitamins such as B and K.3 Some of these nutrients are absorbed directly throughout the wall of the caecum, but most are returned to the rabbit via ingestion of its caecotropes. The caecal contents are moved by periodic contractions through the large bowel, resulting in the production of caecotropes. Caecotropes are soft, green, and mucus coated and have a stronger odour than the hard faeces. They often appear in "grape-like" clusters. Rabbits have a crepuscular life-style and tend to eat a larger volume of food at dawn and dusk.. They produce hard faeces for approximately the first 4 hours after a large meal, with the caecotropes produced during the next 4 hours.3 Rabbits ingest the nutrient-rich caecotropes directly from the anus, so they are rarely seen in a healthy rabbit. The mucous coating protects the living organisms in the caecotrope from the acid pH of the stomach for up to 6 hours. Most of the caecal bacteria are ultimately killed and their contents absorbed, but a few may pass intact into the intestine.6 It is interesting to note that dietary factors such as energy, fibre and protein will affect the quantity of caecotropes consumed. An energy deficiency or high dietary fibre will result in the total intake of caecotropes. Excess of dietary energy or protein may result in incomplete caecophagy, with caecotropes being left on the cage floor. Since caecotropes contain vital nutrients for the health of the rabbit, it is important to make sure that debilitated rabbits have access to them. Rabbits that are obese or have perineal skin flaps or spinal, vestibular, CNS or dental disease are at risk of developing nutritional deficiencies from the inability to ingest caecotropes. Often these rabbits will find and eat their caecotropes if a solid floored cage is used or if the owners place the fresh caecotropes where they can be reached by the pet. Consider vitamin, amino acid and fatty acid supplementation of the diet if a rabbit patient is not producing or ingesting normal caecotropes for a lengthy period. GASTROINTESTINAL DISORDERS Enteric disorders in rabbits can be caused by anything that disrupts normal GIT motility, the pH of the caecum or the stomach, the fluid content of the ingesta (particularly in the hindgut), or the hindgut flora. Some of these factors include low dietary fibre, high dietary starch or protein, inadequate water intake, inactivity, stress, inappropriate antibiotic use, "bulk laxatives" (such as psyllium, which retains fluid in the proximal colon), intestinal parasites, genetic predisposition, and concurrent disease outside of the GIT. Enterotoxaemia The most severe form of enteritis, enterotoxaemia, is caused by a proliferation of Clostridium spiroforme and the resultant iota-like toxin production. The disease is seen most often in newly weaned rabbits (4 to 8 weeks of age) whose GIT has recently been populated with microbes. 7 Enterotoxaemia may also be created by administering inappropriate antibiotics to rabbits at any age, although adult rabbits are more resistant. Antibiotics to avoid include clindamycin, lincomycin, penicillin (orally, although in healthy rabbits on a good diet it may be used parenterally), ampicillin, amoxicillin, amoxicillin-clavulanic acid, cephalosporins and erythromycin.9 True enterotoxaemia results in severe depression, profuse watery diarrhoea often tinged with blood, and death within 24 to 48 hours. It is extremely difficult to treat this disease, and the best course of action is prevention through a healthy diet, low stress environment and avoidance of inappropriate antibiotic use. Chronic Soft Stools By far, the most common form of enteritis seen in the adult pet rabbit is characterised by a chronic soft stool produced intermittently throughout the day. The abnormal stools are mixed with normal hard faeces and may range from "pudding-like" to soft clumps that adhere to the rabbit’s perineum or environment. The soft stools have a strong odour and high mucus content, which identify them as abnormally produced caecotropes. The rabbit may act relatively normally otherwise, although some patients will exhibit periodic anorexia and abdominal discomfort. Many of these patients are on a diet that is either low in fibre and/or high in starch and are overweight and sedentary as well. The cause of this enteritis is likely a mild caecal dysbiosis. Long-term progression of this disease may result in nutritional deficiencies (due to the inability of the rabbit to ingest the liquid caecotropes) and, ultimately, to the more serious condition of GIT ileus. Uncomplicated cases of intermittent soft stools in the rabbit are nearly always treatable through correction of the diet. I have had success feeding these animals on a 100% ad libitum grass hay diet for 2 weeks (or until the stools have returned to normal, which may take up to 2 months in some cases). Once the stools are normal, fresh leafy greens, vegetables and fruits are reintroduced over the next 2 weeks. Commercial pelleted foods and high starch foods (grains, legumes) are never returned to the diet. This diet not only results in a rapid correction of the dysbiosis, but reduces excess weight naturally and safely. Rabbits should also be encouraged to exercise, and fresh water should be available to them at all times. A word of caution before embarking on this therapeutic plan: make sure that the rabbit is familiar with hay and is eating it before removing all other foods from the diet. Intestinal Ileus Another common GIT disorder is intestinal ileus. This is often the result of a chronic GIT hypomotility related to an inappropriate diet as was discussed with gastric stasis. The signs of intestinal ileus are similar to those of gastric stasis (in fact, these disorders often occur together) and include a gradual decrease in appetite and the size and number of hard stools produced. The rabbit eventually becomes anorectic and sits hunched with abdominal pain. Radiography reveals a gas-filled intestinal tract and caecum. Often there is a dense mass of ingesta surrounded by a halo of air in the stomach, and occasionally the same sign is noted in the caecum (caecal impaction). These cases are treated medically as described for gastric stasis. Surgery should be avoided, and mild to moderate cases usually exhibit a return of GIT motility and stool production within 72 hours. Gastrointestinal Obstruction Acute gastric or intestinal obstruction is seen more rarely than gastric stasis or intestinal ileus. Obstructive disease is often confused clinically with ileus. The most common site of obstruction is the pylorus or upper duodenum. The second most common site is the ileocaecocolonic junction (the result being a caecal obstruction). GIT obstructions can also be caused by postsurgical abdominal adhesions. The material that most frequently causes an obstruction is a piece of dried ingesta mixed with hair. This is not a true hairball as is seen in the cat or the ferret, but rather a matrix of food and hair. Other foreign bodies include carpet fibres, plastic or rubber. I have encountered several rabbits that developed complete GIT obstruction after ingesting large amounts of clay cat litter. An accurate history reveals that the rabbit was perfectly normal one day and acutely depressed and anorectic with no stool production the next; in contrast, ileus is characterised by a gradual onset of signs. Radiographically, distention of the GIT with fluid and air is isolated to the stomach and the duodenum with upper GIT obstruction. The stomach of an obstructed patient appears grossly distended with fluid and some air, as opposed to the distinct halo of air seen around the dense, distinct food ball of the gastric stasis patient. If the caecum is obstructed, it will be greatly distended with fluid. GIT obstruction is an emergency situation requiring immediate surgery. It can be very difficult to decompress the stomach via a stomach tube because ingesta readily clog the tube. Administer glucocorticoids for shock, intravenous or intraosseous fluid therapy, and analgesics and provide other supportive care as needed. The prognosis is guarded to poor, even with aggressive therapy. Bacterial Enteritis Bacterial enteritis is rare in the adult rabbit, primarily because of the high stomach pH, which sterilises the ingesta. It is more common in young rabbits, rabbits on a poor diet, or those receiving inappropriate antibiotics for reasons already described. Bacterial enteritis can be caused by infection with E. coli, Clostridium piliforme (formerly Bacillus piliformis; also known as Tyzzer’s disease), Salmonella, Pseudomonas, and Campylobacter-like species.6 Viruses are not a common cause of GIT disease in the pet rabbit in the United States. Rabbit enteric coronavirus and rotavirus have been encountered in commercial rabbitries.6 Rabbit viral haemorrhagic disease is found in China, Europe and Mexico but has not yet entered the United States. Parasitic Disorders Parasitic disorders of the GIT of the rabbit involve coccidia (Eimeria spp.), Cryptosporidium, Passalurus ambiguus, Passalurus nonanulatus, Obeliscoides cuniculi and a variety of cestodes and trematodes.6 Coccidia are the most commonly encountered GIT parasites and may infect either the liver (specifically Eimeria stiedae) or the intestine. Hepatic infections may be subclinical or result in unthriftiness and death. Intestinal infection is often subclinical but occasionally causes severe dehydration and death. Mild infections in healthy rabbits ultimately result in long-term immunity. Treatment for coccidia involves the use of sulfa drugs. One suggested treatment regimen for coccidia is sulfadimethoxine (15 mg/kg PO q12h for 10 days) combined with a trimethoprim-sulfa combination (30 mg/kg PO q12h for 10 days).6 DIET RECOMMENDATIONS The diet I recommend for the noncommercial, non-breeding pet rabbit is ad libitum grass hay and large amounts of fresh leafy greens with added vegetables and fruits. Hay is the most important part of the diet as it provides a naturally high indigestible fibre source and encourages proper tooth wear. Good quality grass hay is also rich in other needed nutrients such as protein, vitamins, minerals and carbohydrates. Grass hays available in the United States include timothy, oat, barley and bromegrass or mixed grasses. Alfalfa hay can be used if grass hay is not available, but it is higher in protein, calories and calcium, which may be undesirable in some patients. I have found it more difficult to return overweight animals to their proper weight and to control some cases of chronic soft stools if patients are fed large amounts of alfalfa hay. In addition, it is not helpful to mix grass and alfalfa hay, as the more palatable alfalfa hay will be eaten exclusively and the grass hay left behind. I have placed pet rabbits on an all hay diet for up to 4 months to treat GIT disorders without ill effect. The three main drawbacks to hay are: Some humans are allergic to hay dust and cannot be exposed to it. · [US only] Hay can contain the eggs of Baylisascaris procyonis (the raccoon roundworm) if it was taken from the top of the pile in an open hay loft or left outside on the ground uncovered. It may be difficult to obtain quality grass hay year-round, making client compliance difficult. A less desirable but usable alternative to loose hay is pressed hay cubes which can be purchased in some pet and feed stores. The second most important part of the diet of the pet rabbit is clean, fresh leafy greens, including kale, collards, beet tops, carrot tops, parsley, dandelion greens, chicory, mustard greens, romaine lettuce, raspberry leaves, basil, mint, endive, radicchio, wheat grass and Swiss chard to name a few. A minimum of three different greens should be fed daily, but the more variety the better. If the rabbit has never had greens, establish it on hay first; then introduce a variety of greens over a few weeks at a minimum amount of 1 cup of tightly packed greens per 1kg body weight per day. Once rabbits are well established on this diet, the total volume of greens is unrestricted. Rarely, an owner will observe a soft stool in relation to a particular green food (parsley is a common culprit); that food may be removed from the diet. Rabbits that ingest large amounts of greens will greatly decrease the amount of water they drink. Other fresh foods fed in smaller amounts include carrots, pea pods, green pepper, broccoli, Brussels sprouts, melons, berries, apples, pears, peaches, papaya, pineapple and mango. Avoid feeding legumes or grains (including bread, cookies, oats and corn), which are high in starch, and grapes and bananas, which often are "addictive". Fresh food provides not only important fibre, fluid and other nutrients but also likely provides stimulation in terms of different tastes and textures of foods. I have found that rabbits fed a diet comprised primarily of grass hay and fresh foods are less likely to chew their own or other rabbits’ fur and are less likely to spend time chewing on cage bars, dry wall, furniture and newspaper. Owners like to feed "treat" foods, and they can be encouraged to find special fresh foods, particularly fruits, that can be used in this manner. In addition, fresh non-toxic tree branches, blocks of untreated soft wood and unfinished grass or wicker baskets can all serve as excellent "dietary toys" as well as exercise for the teeth. The least important part of the diet of the noncommercial, nonreproductive pet rabbit is commercial pellets. In my experience the ad libitum feeding of a variety of commercial pellets has contributed to innumerable cases of dental disease, obesity, enteritis, ileus and gastric stasis. Since removing or severely restricting commercial pellets in the diet of the pet rabbit and supplementing it with hay and greens, the frequency with which these conditions are seen has been reduced dramatically. Commercial pellets are based on alfalfa meal, grains and other supplements and are designed for the rapid growth of the commercial or laboratory rabbit that is usually not intended to live out its full life span. To the credit of some feed companies there are commercial pellets now being produced that are more suitable for the pet rabbit. They have a high fibre content (18% minimum), a moderate to low protein level (14%-16%) and low calcium and fat. These "pet rabbit" pellets can be used in situations where a weight gain is desired, where fresh hay cannot be fed, or where client compliance feeding the "natural" diet is poor. Unless a weight gain is desired, the amount of any pellets used in the pet rabbit diet should be restricted. Fresh water is offered free choice and should not be contaminated with supplements, which may reduce water consumption. As mentioned, rabbits on a diet high in fresh foods drink considerably less water than a rabbit on a pelleted or all hay diet. Rabbits should be encouraged to exercise daily; ideally, they should not be kept in cages bur rather in large pens. Daily exercise has multiple health benefits including improvement of GIT motility. The solution to many rabbit health problems is feeding an appropriate diet for this species. The biggest obstacle to a healthy diet is client resistance to abandoning what has historically been touted as the "proper" rabbit diet (i.e. pellets). In addition, clients like to feed their rabbits an assortment of treat foods that are usually high in fat or starch, which only complicates matters further. It is important to remember that often they neglect to mention these treat foods when you are trying to resolve a problem case (e.g. chronic soft stools). However, with aggressive education efforts and demonstration of the immense benefits of a proper diet, clients can be won over. Once a lethargic, overweight rabbit that produced soft stools has been on a healthy diet for a few weeks, I often hear comments like "I didn’t know he had that much energy! He isn’t a ‘couch rabbit’ any more". References 1 Krull WH: Coprophagy in the Wild Rabbit (Sylvilagus nutalli granger) (Allen). Vet Med 35:481483, 1954. 2 Crossley DA: Dentistry for Small Animals other than Cats and Dogs. Second Annual Midwest Exotic Pet Seminars Proceedings, Schaumburg, IL, March 1997. 3 Cheeke PR: Rabbit Feeding and Nutrition. Orlando, Academic Press, 1987, pp 20-21. 4 Smith HW: The Development of Flora of the Alimentary Tract in Young Animals. J Pathol Bacteriol 89:95-122, 1965. 5 Brooks DL: Rabbit Gastrointestinal Disorders, in Kirk RW (ed): Current Veterinary Therapy VIII. Philadelphia, WB Saunders, 1983, pp 654-657. 6 Brooks DL: Nutrition and Gastrointestinal Tract Physiology, in Hillyer EV, Quesenberry KE (eds): Ferrets, Rabbits, and Rodents: Clinical Medicine and Surgery. Philadelphia, WB Saunders, 1977, p 148. 7 Harkness JE, Wagner JE: The Biology and Medicine of Rabbits and Rodents, ed 4. Media, PA, Williams & Wilkins, 1995 8 Forsyth SJ, Parker DS: Nitrogen Metabolism by the Microbial Flora of the Rabbit Caecum. J Appl Bacteriol 58:363-369, 1985. 9 Jenkins JR: Gastrointestinal Disorders, in Hillyer EV, Quesenberry KE (eds): Ferrets, Rabbits, and Rodents: Clinical Medicine and Surgery. Philadelphia, WB Saunders, 1977, pp 176-188. © Waltham USA, Inc.