The digestive tract in horses is a complex system responsible for the breakdown and absorption of nutrients essential for maintaining equine health. It includes several key components: the mouth, esophagus, stomach, small intestine, cecum, large colon, small colon, and rectum. Each segment of the digestive tract performs specific functions, from the mechanical breakdown of feed to the enzymatic digestion and absorption of nutrients. The horse's digestive system is uniquely adapted to process fibrous plant material, relying heavily on microbial fermentation, particularly in the hindgut. This page compiles peer-reviewed research studies and scholarly articles that explore the anatomy, physiology, and function of the equine digestive tract, as well as common disorders and their management.
Cymbaluk NF, Schryver HF, Hintz HF, Smith DF, Lowe JE.The effects of molybdenum ingestion on copper metabolism were investigated in bile duct-cannulated ponies fed diets containing 1.01, 27.4 and 107.3 ppm molybdenum. Copper metabolism was assessed by examination of 64Cu and stable copper excretion in feces, bile and urine. The addition of molybdenum to the diet decreased copper absorption and retention as a consequence of increased excretion of dietary copper in feces and increased excretion of absorbed copper in bile. Only 5% or less of the absorbed copper was excreted in urine regardless of dietary treatment. Biliary molybdenum excretion was o...
Minder HP, Merritt AM, Chalupa W.Feces from 13 healthy horses and 8 horses with chronic diarrhea were subjected to an in vetro fermentation procedure that had been developed for rumen fluid. Fermentations were conducted over 6 hours in a closed system, with and without an essential amino acid (EAA) mixture being added to the basic starch-buffer medium. The addition of EAA caused no significant difference in results of fermentation of feces from healthy horses. For diarrheic animals, there was a significant (P less than 0.01) increase in gas and total volatile fatty acids production whether EAA were present or not, and alpha-a...
Clayton HM, Duncan JL, Dargie JD.The mechanisms involved in the pathophysiological disturbances associated with the presence of mature Parascaris equorum in the small intestine were investigated with radioisotopic techniques. The results suggested that, compared with worm-free controls, infected foals had a reduction in gut motility, an increase in the body solids ratio, a lowering of the body pool of albumin and a decreased ability to incorporate dietary methionine into plasma protein.
Wolter R, Nouwakpo F, Durix A.A complete pelleted diet (table 1) containing 11 p. 100 of crude protein and 17 p. 100 of cellulose (ADF-lignine according to the method of Van Soest) was distributed to ponies and rabbits. Total digestibilities, partial cumulative digestibilities in different compartments of the digestive tract (stomach, small intestine, caecum, proximal and distal colon) and changes in biochemical composition, measured by pH, VFA, lactic acid and ammonia concentrations, were compared. Total digestibilities (table 2). There was no significant difference in dry matter between the two species. Crude protein dig...
Tisserand JL, Boulard F, Deponge P.We studied a caecum-cannulated pony fed hay libitum in order to determine the mineral elements likely to cause deficiencies which decrease microbial activity in the caecum. During four successive 1-month periods, we observed the effects of adding 50 mg of iron, 1 mg of cobalt, 40 mg of copper and 200 mg of zinc, respectively. Except for the introduction of zinc, which greatly diminishes cellulolytic activity measured with the nylon bag technique, there was no significant modification in that activity as shown by pH, NH3-nitrogen, total nitrogen and volatile fatty acid production (table 1). It ...
Tisserand JL, Ottin Pecchio M, Rollin G.The cellulolytic activity in the large intestine of the pony varies according to the form and the composition of the feed. This activity was measured on two caecal and ventral colon-cannulated ponies receiving the following 4 diets during four successive 6-week periods: --6 kg of hay, --4 kg of hay + 1 kg of oats, --6 kg of ground, pelleted hay. --5 kg of a ground, pelleted blend of 80 p. 100 hay and 20 p. 100 oats. Adding oats to a hay feed increased the cellulolytical activity in the caecum and the colon, whereas grinding and pelleting hay alone or hay enriched with oats diminished that acti...
Hjortkjaer RK.In the first series of experiments enema containing a water soluble marker was given to two groups of horses fasted for 1 and 18 hours, respectively. The marker was only in 1 out of 14 experiments found orally of the pelvic flexure (Table II). Fasting seemed to facilitate the flow in oral direction. The results were discussed with the possible significance in the treatment of large intestinal impactions. In the second series the effect of enemas on water, electrolyte and acid-base status in the acute furosemide-dehydrated horse was studied. Moderate acidifying and hypokalemic effect was encoun...
Sellers AF, Lowe JE, Brondum J.The equine large colon is the major cellulose-fermentation locus of these species. The area of juncture of the ventral and dorsal divisions of the equine large colon was characterized, in 13 chronic unanesthetized animals and in 25 in vitro preparations, as an area of resistance to aboral flow. This is a probable pacemaker area. The reservoir function of this part of the colon is apparently facilitated by the presence of a pacemaker at this level. There was a general rise in recorded intraluminal pressure peaks during the first hour of feeding. This characteristic was not altered after resecti...
Steffey EP, Johnson BH, Eger EI, Howland D.Breathing 79% nitrous oxide (N2O) in oxygen increased the rate of accumulation of bowel gas during intraluminal bowel segment infusions of hydrogen, methane (CH4), air, or carbon dioxide (CO2) in four pentobarbital-anesthetized dogs more than did breathing 100% oxygen. A N2O-associated increase in the volume of naturally produced intestinal gas in five halothane-anesthetized ponies corroborated the findings in the dog studies. In a second group of four dogs a bolus of CH4 or CO2 was injected into the bowel lumen. When the dogs breathed O2 the bowel gas volume decreased. Gas was virtually absen...
Irwin DH, Howell DW."Cube colic" is described and vain attempts to cause the problem on an experimental basis are recorded. It is concluded that not all horses are susceptible to cube colic and if colic occurs in horses being fed on cubes, it is not necessarily due to feeding of cubes. Nomenclature of bowel disorders is considered.
Bayley HS.The research article explores the comparative physiology of the hindgut in different animals, its key processes, and its implications on nutrition. It underscores the significance of gut microorganisms in breakdown […]
Willard JG, Willard JC, Wolfram SA, Baker JP.Three cecal-fistulated horses were used in a 3 × 3 latin square experiment to determine the influence of diet and of cecal infusions of Na2C03 on cecal fermentation and feeding behavior. The three treatments were hay, concentrate and concentrate plus hourly infusions of Na2CO3. Cecal fluid samples and cecal pH readings were taken at zero through 11 hr following feeding at the end of each experimental period, and animal activity was measured by the use of a movie camera set to take 5 sec of film every 5 minutes. Cecal pH was significantly lower at 4, 5 and 6 hr following feeding for the horses...
La CelluleJanuary 1, 1977
Volume 72, Issue 1-2 5-25
Meuleman A, Latteur B.The topography of Cochliatoxum periachtum is studied. The cuticle is very thick. The digestive, nuclear and excretory systems are located in the ectoplasm, they are not surrounded by the mesoplasm. The endoplasm is very large, it contains vegetable inclusions. By the features of its topography, this species is similar to some species belonging to the Family Ophryoscolescidae and, especially, to Eudiplodinium giganteum.
Reinertson EL.Thirteen horses and ponies were used to evaluate an anastomotic technique utilizing a modified Gambee suture pattern. The modified Gambee technique was compared with a single-row everting technique and a double-row inverting technique. The double-row inverting technique predisposed to intestinal obstruction and resulted in minimal adhesions. The everting technique predisposed to leakage of intestinal contents and subsequent peritonitis. Adhesions were formed at the site of the anastomosis to such an extent as to jeopardize the proper position of the bowel. The technique did not cause any lumin...
Barlet JP, Davicco MJ, Coxam V.Intestinal absorption of inorganic phosphorus. In most mammalian species inorganic phosphorus (P) is absorbed at the duodenal and jejunal level. However in horses some P is absorbed from the large intestine, whereas in ruminant animals some absorption also takes place in the forestomachs. The structure of the putative phosphate-sodium carrier of the brush border from enterocytes still awaits identification. P absorption is modulated both by endocrine (calcitriol, triiodothyronine) and nutritional factors (minerals of the diet, chemical form of phosphorus). Regulation of salivary P secretion, i...
Ralston SL, Freeman DE, Baile CA.The roles of volatile fatty acids (VFA) and of the large intestine in the control of feeding in ponies were investigated. Ponies with cecal fistulas were adapted to ad libitum access to pelleted feed. Treatment solutions were given as a bolus 15 min before the animals were allowed free access to feed after a 4-h fast. Each dose of VFA solution was tested in a crossover design with a water control. When the ponies were permitted to eat after the treatments, the latency to eat, first meal size, and duration and first intermeal interval were recorded. Feed intakes were measured at 3 and 18 h afte...
Bertone AL, Toofanian F, Stashak TS.Lactase, maltase, sucrase, and alkaline phosphatase activities were determined in the intestinal mucosa from 3 locations in the small intestine and 4 locations in the large intestine 1 year after extensive large-colon resection (group 1; n = 5) and 1 year after sham operation (group 2; n = 3) in horses. Lactase, maltase, and sucrase activities were similar (P greater than 0.05) between group-1 and group-2 horses in all locations measured in the intestinal tract. Alkaline phosphatase activity in the remaining large colon of group-1 horses was significantly (P less than 0.05) greater than the ac...
Mobarak MS, Ryan MF.Light, scanning, and transmission electron microscopy were employed to provide further data on the putative origins of the immunogenic secretory-excretory product (ESP) of Strongylus vulgaris (Looss 1900). The sharply delineated but superficial attachment to the equine caecum by the mouth leaves behind an oval area devoid of epithelial cells. Attachment does not extend deeply enough to reach the muscularis mucosa layer of the equine intestine. The progressive digestion of the ingested plug of tissue (epithelial cells, blood cells and mucous) was visualized. The coelomocytes, floating cells and...
Williams TL, Mochal-King C, Rude BJ, Weed BC, Nicodemus MC.Cecal and ileal cannulations have previously been used to determine equine digestibility and define gastrointestinal physiology. However, research has been limited to specific portions of the equine digestive tract lacking utilization of dual cannulations. The purpose of this study was to develop and implement surgical and preoperative and postoperative procedures for cannulation of dual sections, ileum and cecum, of the equine digestive tract including development of recommendations for addressing postsurgery complications. A total of 8 ponies went through surgical procedures for the placemen...
Wawrzeńska M.In this paper, the effect of ascorbic acid on motoric activity of the intestines of rabbits, pigs, cows, sheep and horses has been determined, and a possible participation of the adrenogenic system in this mechanism has been shown. In experiments in vitro the motility of the duodeum, jejunum, caecum and colon in the animals mentioned abowe was recorded by the method of Magnus after administration of ascorbic acid. Diastolic reactions were observed in all animals, which were much greater in small intestines than in large ones. To elucidate the diastolic mechanism under the influence of ascorbic...
Fowler AL, Pyles MB, Hayes SH, Crum AD, Lawrence LM.Across the equine literature, estimates of true P digestibility range from -23% to 79%. This large range cannot be explained by differences in P intake or phytate-P intake alone. However, differences in endogenous P secretion into the GI tract may explain the variation. In horses, excess absorbed P is not excreted in the urine but is re-secreted into the GI tract, increasing faecal P and leading to estimates of low P digestibility. Thus, accurate estimates of P digestibility can only be obtained if absorbed P is retained in the horse. The objective of this study was to examine P digestibility ...
Reed L, MacNicol JL, Charchoglyan A, Brewer D, Murrant C, Pearson W.The objective of this study was to evaluate the effect of a botanical-based equine nutraceutical on contractility of gastric smooth muscle in vitro. Gastric ulcers are prevalent in performance horses and negatively impact horse welfare. Gastric hypermotility has been positively associated with the development of gastric ulceration in nonequine species, and reduction of hypermotility may be protective against their development. Stomachs from 12 pigs processed for food at a provincially inspected abattoir were collected within 1 hour of slaughter. Explants of nonglandular gastric tissue were p...
Cottrell DF, Jones AF, Potter KE.Electromanometric recordings of caecal pressures were made in chronically cannulated ponies. Endoscopy allowed direct observation of the caecal base and cupola while recording intracaecal pressures and enabled a functional interpretation of caecal pressure profiles. Using gas-tight seals, the caecal gas cap baseline pressure (tone) was usually below atmosphere, becoming less negative postprandially. Diurnal variations were observed in the amplitude and frequency of caecal pressure profiles and the direction of caecal movements, during which intra-caecal gas cap pressures briefly became positiv...
Tisserand JL, Boulard F, Deponge P.We studied a caecum-cannulated pony fed hay libitum in order to determine the mineral elements likely to cause deficiencies which decrease microbial activity in the caecum. During four successive 1-month periods, we observed the effects of adding 50 mg of iron, 1 mg of cobalt, 40 mg of copper and 200 mg of zinc, respectively. Except for the introduction of zinc, which greatly diminishes cellulolytic activity measured with the nylon bag technique, there was no significant modification in that activity as shown by pH, NH3-nitrogen, total nitrogen and volatile fatty acid production (table 1). It ...
Decloedt AI, Bailly-Chouriberry L, Vanden Bussche J, Garcia P, Popot MA, Bonnaire Y, Vanhaecke L.Traditionally, steroids other than testosterone are considered to be synthetic, anabolic steroids. Nevertheless, in stallions, it has been shown that β-Bol can originate from naturally present testosterone. Other precursors, including phytosterols from feed, have been put forward to explain the prevalence of low levels of steroids (including β-Bol and ADD) in urine of mares and geldings. However, the possible biotransformation and identification of the precursors has thus far not been investigated in horses. To study the possible endogenous digestive transformation, in vitro simulations of t...
Bennett DG.Routine examination of the digestive system relevant to purchase includes gross examination of the feces, auscultation of the abdomen, and examination of the oral cavity and teeth for verification of age and identification of any gross abnormality. Further examination is indicated only when there is a history of diseases such as colic, especially colic surgery, or esophageal obstruction. Special procedures that might be indicated, depending upon the history and findings on the routine physical examination, include rectal palpation of the abdominal viscera; endoscopic examination of the esophag...
Frape DL.The majority of recent experiments in equine nutrition relates to the metabolism of protein, non-protein nitrogen, carbohydrate, calcium and phosphorus. An understanding of the significance of the nitrogen cycle is emerging in which there is a two-way movement of compounds across the wall of the G.I. tract. Some quantitative estimates of the role of microflora in this process have been made for comparison with similar processes in ruminants. However, the pathway by which the microflora contributes to the nitrogen economy and the extent to which this occurs is not established in the horse. It i...
Reinemeyer CR, Herd RP.Haustral portions of intestine of 6 horses were isolated by excising the taeniae coli from the cecum and the ventral colon. Uniform 5-cm X 5-cm sections were cut from the haustra and were illuminated from the serosal side with a strong light source (mural transillumination). Cyathostome larvae encysted in the mucosa and submucosa were observed at 15 X magnification and counted. Two separate counts of the larvae in 80 replicates of tissue by the mural transillumination technique (MTT) revealed no significant (P less than 0.05) difference between sample means. Larvae in tissue sections were coun...
Wolford AN, Coverdale JA, Leatherwood JL, Pinchak WE, Anderson RC, Wickersham TA.Eight previously cecally cannulated Quarter Horse geldings were utilized in a crossover design with two 28-d periods with a 21-d washout period between to evaluate the influence of housing on the cecal environment and dry matter intake (DMI). Horses were adapted to diet and housing from day 1 to 19, DMI was determined from day 20 to 24, and cecal fluid was collected on day 28. Horses were paired by age and body weight (BW) and randomly assigned to treatment. Treatments consisted of housing horses individually in stalls or group housed in a pen. Regardless of treatment, all horses were individu...
Tisserand JL, Ottin Pecchio M, Rollin G.The cellulolytic activity in the large intestine of the pony varies according to the form and the composition of the feed. This activity was measured on two caecal and ventral colon-cannulated ponies receiving the following 4 diets during four successive 6-week periods: --6 kg of hay, --4 kg of hay + 1 kg of oats, --6 kg of ground, pelleted hay. --5 kg of a ground, pelleted blend of 80 p. 100 hay and 20 p. 100 oats. Adding oats to a hay feed increased the cellulolytical activity in the caecum and the colon, whereas grinding and pelleting hay alone or hay enriched with oats diminished that acti...
Kawaida MY, Maas KR, Moore TE, Reiter AS, Tillquist NM, Reed SA.To determine the effects of astaxanthin (ASTX) supplementation on the equine gut microbiota during a deconditioning-reconditioning cycle, 12 polo ponies were assigned to a control (CON; n = 6) or supplemented (ASTX; 75 mg ASTX daily orally; n = 6) group. All horses underwent a 16-week deconditioning period, with no forced exercise, followed by a 16-week reconditioning program where physical activity gradually increased. Fecal samples were obtained at the beginning of the study (Baseline), after deconditioning (PostDecon), after reconditioning (PostRecon), and 16 weeks after the ces...
Silva AH, Lucena JEC, Taran FMP, Cunha DS, Almeida JAT, Lima RS, Dias WS, Santiago JM.This study aimed to compare the digestibility of tropical grasses by horses by the in vivo method using mobile nylon bags with the in vitro digestibility method using horse feces as a source of inoculum. Five horses were used in a 2 × 5 factorial design with randomized blocks featuring two methods (in vivo and in vitro) and five grasses: Tifton 85 hay (Cynodon spp.), sixweeks threeawn grass (Aristida adsencionis, Linn), Alexandergrass (Brachiaria plantaginea (Link) Hitchc.), capim-de-raiz (Chloris orthonoton, Doell), and Sabi grass (Urochloa mosambicensis). No difference (P>0.05) was foun...
Elisabeth Müller C.A study comprising 74 colic and 74 control horses admitted to an animal hospital was performed. Faecal samples were collected and analysed for dry matter concentration, particle size distribution using wet-sieving, and sand presence through a sand sedimentation test. Data on horse breed, age, gender and basic feeding variables was collected and analysed using χ-tests. Faecal dry matter concentration, particle size distribution and sand score was compared between colic and non-colic horses, and between horses with different colic types, using one-way ANOVA. Results showed that colic and non-co...
Barton CK, Hector RC, Hendrickson DA, Kawcak CE, Nelson BB, Goodrich LR.To compare 3 perioperative feeding regimens and their effect on anesthetic complications, manure output, and colic proportion in healthy horses. Methods: 45 horses presenting for elective orthopedic procedures were randomly assigned to 1 of 3 groups: not fasted (NF; continuous access to hay perioperatively), fasted muzzled (FM; 10-hour preoperative fast with slow refeeding postoperatively and muzzle placement), or fasted not muzzled (FNM; same as FM without muzzle placement). Anesthetic protocol was standardized. Outcomes compared between groups included anesthesia time, arterial oxygenation, ...
Safaee Firouzabadi MS, Paidar Ardakani A.Many studies have shown a strong correlation between Hindgut Acidosis (HGA) and the occurrence of laminitis in horses; therefore, the early diagnosis of HGA is essential. In this study, we investigated changes in the plasma concentrations of lipopolysaccharide-binding protein (LBP) and serum amyloid A (SAA) as inflammatory markers in horses with laminitis. Sixteen healthy male Arabian horses that had cecal cannulation without visible laminitis or general symptoms were randomly divided into two groups. The horses were fed two different diets in a forage-to-concentrate ratio. Blood samples were ...
Azizi S, Mazaheri-Khameneh R, Davoodi F, Gooran MM, Ghoreishi SS, Mozaffari N, Dalir-Naghadeh B.In this case report, we present an unusual clockwise torsion of left colon around mesenteric root in a 10-month-old Arab filly, highlighting the clinical presentation, diagnostic approach and successful surgical intervention. A 10-month-old Arab filly weighing approximately 250 kg was referred with signs of acute abdominal pain. The history revealed anorexia, restlessness and severe abdominal pain that had begun the previous day. The local practitioner had previously administered flunixin meglumine, an analgesic, but it proved ineffective in relieving the pain. Upon physical examination, the ...
Bachmann M, Schusser GF, Wensch-Dorendorf M, Pisch C, Bochnia M, Santo MM, Netzker H, Woitow G, Thielebein J, Kesting S, Riehl G, Greef JM....Concentrations of starch, mono- and disaccharides, fructans, hemicellulose and cellulose were analysed in feed and gastric digesta of horses in relation to acid insoluble ash as a marker indigestible in the stomach. Twenty-four horses were allocated to pasture 24 h/d (PST; n = 4), hay ad libitum (HAY; n = 8), hay ad lib. and oats at 1 g starch/kg body weight (BWT)/meal (OS1; n = 6) and hay ad lib. and oats at 2 g starch/kg BWT/meal (OS2; n = 5). One horse was excluded from the analysis. The horses were fed the ration a minimum of 34 days. Following euthanasia and dissection, digesta was sample...
Sibthorpe PEM, Fitzgerald DM, Sillence MN, de Laat MA.Equine insulin dysregulation (ID) is a significant metabolic problem because the hyperinsulinaemia that develops increases the animal's risk of developing laminitis, a debilitating foot condition. The role of gastrointestinal factors, such as incretin hormones, in the pathogenesis of ID and hyperinsulinaemia in horses is poorly understood, particularly in comparison to other species. Glucagon-like peptide-2 (GLP-2) is an intestinotrophic peptide released from L cells in the gastrointestinal tract and is implicated in metabolic dysfunction in other species. The aim of this study in vitro was to...
St Blanc MP, Mirza MH, Riggs LM, Leise BS.The aim of the study was to evaluate time to closure, bursting pressures and luminal diameters of a single and double-layer end-to-end anastomoses in normal equine descending colon. Methods: Experimental study. Methods: Eight adult horses. Methods: Four segments of descending colon from each horse were randomly assigned to a control (n = 8, CON), one-layer (n = 12, group 1; continuous Lembert pattern), or two-layer group (n = 12, group 2; simple continuous oversewn with Cushing pattern). Anastomoses were performed, and time to closure and luminal diameter were measured. Bursting pr...
Daniels SP, Whiteside EJ, Martin S, Moore-Colyer MJS, Harris P.Feedstuffs, especially ensiled forage, may be detrimental to equid oral health by exposing the oral cavity to low pH. This study aimed to identify if salivary pH was altered by 1) a range of different feedstuffs and (2) specifically by feeding haylages with differing nutrient profiles. Two studies were conducted. The first measured oral pH following five feedstuffs, (hay, haylage, unmolassed sugarbeet pulp, unmolassed alfalfa chaff and rolled oats), tested individually over five days. Saliva (≥1 ml) was collected in triplicate, prior to feeding, directly after ingesting 500 g of each feedstu...
Kopec EK, Stevens M, Crowe O, Wright C, Suthers J.The aim of the present study was to investigate the long-term survival and postoperative complications in horses that had undergone small intestinal resection and a modified anastomosis technique using a partially stapled, functional end-to-end anastomosis. Methods: Observational retrospective case series. Methods: A total of 25 horses underwent small intestinal resection and a partially stapled, functional end-to-end anastomosis. Methods: A modified technique to create a partially stapled, functional end-to-end anastomosis is described. The clinical case records of all horses that fulfilled t...
Barton CK, True A, Fenn M, Thampi P, Watson A, Smanik LE.The objective of this Case Report was to describe the clinical manifestation and postmortem findings of a horse evaluated for recurrent colic diagnosed with pyloric aplasia and an ectopic duodenal papilla. Methods: 13-year-old Quarter Horse gelding. Unassigned: The horse was presented to a referral center for signs of colic that were unresponsive to management on the farm. No abnormalities were noted on rectal palpation; however, upon passage of a nasogastric tube, 31 L of net gastric reflux was obtained. Gastroscopy noted grade IV squamous ulceration and moderate glandular ulceration, severe ...
Lopes MAF, Hollenbach E, Schliewert EC, Murphy MA, Corrêa F.A mare was admitted for progressive swelling of the neck, lethargy and anorexia. Radiography revealed perilaryngeal, peritracheal, and periesophageal emphysema. Endoscopy revealed pharyngeal and tracheal roof collapse, and a small laceration on the trachea. Treatment with antimicrobials, fluids, and flunixin was initiated. To prevent exacerbation of the emphysema, temporary tracheostomy was performed. Clipping for the tracheostomy revealed a hoofprint mark on the ventral neck. Subsequent endoscopies revealed laceration of the esophagus and guttural pouch septum communicating with the visceral ...
Medina-Bolívar AL, Faleiros RR, Martínez-Aranzales JR.Limited information is available on gastric diseases in neonatal foals as compared with extensive studies in young, adult, and geriatric horses. Reports on fetuses are scarce. Objective: Assess at necropsy stomachs of horse, donkey and mule fetuses in the third trimester of gestation to characterize lesions present during intrauterine life. Methods: Forty-six fetal stomachs from both sexes (21 horses, 21 donkeys, and 4 mules) in the third trimester of gestation were collected from a processing plant immediately after slaughter. Methods: Measurements of longitudinal and transverse axes, weight ...
