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Topic:Physiology

The physiology of horses encompasses the study of the biological functions and processes that occur within the equine body. This includes the examination of various systems such as the cardiovascular, respiratory, musculoskeletal, digestive, and nervous systems. Understanding equine physiology is essential for comprehending how horses adapt to different environmental conditions, perform physical activities, and respond to health challenges. Research in this field often focuses on the mechanisms of energy metabolism, thermoregulation, and muscle function during exercise, as well as the physiological responses to stress and disease. This page compiles peer-reviewed research studies and scholarly articles that explore the diverse aspects of equine physiology, providing insights into the biological processes that support the health and performance of horses.
Equine herpesvirus type 1 (EHV-1) utilizes microtubules, dynein, and ROCK1 to productively infect cells.
Veterinary microbiology    August 8, 2009   Volume 141, Issue 1-2 12-21 doi: 10.1016/j.vetmic.2009.07.035
Frampton AR, Uchida H, von Einem J, Goins WF, Grandi P, Cohen JB, Osterrieder N, Glorioso JC.To initiate infection, equine herpesvirus type 1 (EHV-1) attaches to heparan sulfate on cell surfaces and then interacts with a putative glycoprotein D receptor(s). After attachment, virus entry occurs either by direct fusion of the virus envelope with the plasma membrane or via endocytosis followed by fusion between the virus envelope and an endosomal membrane. Upon fusion, de-enveloped virus particles are deposited into the cytoplasm and travel to the nucleus for viral replication. In this report, we examined the mechanism of EHV-1 intracellular trafficking and investigated the ability of EH...
Does the microbial flora in the ejaculate affect the freezeability of stallion sperm?
Reproduction in domestic animals = Zuchthygiene    August 6, 2009   Volume 44, Issue 3 518-522 doi: 10.1111/j.1439-0531.2008.01267.x
Ortega-Ferrusola C, González-Fernández L, Muriel A, Macías-García B, Rodríguez-Martínez H, Tapia JA, Alonso JM, Peña FJ.In an attempt to evaluate the possible relationship between the microbial flora in the stallion ejaculate and its ability to freeze,three ejaculates from five stallions were frozen using a standard protocol. Before freezing, an aliquot was removed for bacteriological analysis. Bacterial growth was observed in all the ejaculates studied. The isolated microorganisms were:Staphylococcus spp. and Micrococcus spp. (in all the stallions), beta-haemolytic Streptococcus (in stallions 3 and 4), Corynebacterium spp. (in stallions 1, 3-5), Rhodococcus spp. (in stallion number 2), Pseudomonas spp. (in sta...
CE provides evidence of the stereoselective hydroxylation of norketamine in equines.
Electrophoresis    August 5, 2009   Volume 30, Issue 16 2912-2921 doi: 10.1002/elps.200900221
Schmitz A, Theurillat R, Lassahn PG, Mevissen M, Thormann W.CE with multiple isomer sulfated-CD as selector was used for the simultaneous analysis of the stereoisomers of ketamine, norketamine, 5,6-dehydronorketamine and hydroxylated metabolites of norketamine in liquid/liquid extracts of (i) in vitro incubations with ketamine or norketamine and equine liver microsomes and (ii) plasma and urine of ponies receiving a target-controlled infusion of ketamine under isoflurane anesthesia. Hydroxynorketamine metabolites with the hydroxy group at the cyclohexanone ring could be shown to be formed stereoselectively both in vitro and in vivo. Due to the lack of ...
Characterization of bradykinin-induced endothelium-independent contraction in equine basilar artery.
Journal of veterinary pharmacology and therapeutics    August 4, 2009   Volume 32, Issue 3 264-270 doi: 10.1111/j.1365-2885.2008.01037.x
Ueno D, Yabuki A, Obi T, Shiraishi M, Nishio A, Miyamoto A.We investigated the effect of bradykinin (BK) on isolated equine basilar arterial rings with and without endothelium. BK induced concentration-dependent contraction of resting arterial rings and no relaxation when the rings were precontracted by prostaglandin F(2alpha). The maximal response and pD(2) value were 161.2 +/- 28.1% (to 60 mm KCl-induced contraction) and 8.24 +/- 0.25 respectively. The cumulative concentration-response curve for BK was not shifted to the right by des-Arg(9)-[Leu(8)]-BK (a B(1)-receptor antagonist), HOE140 (a B(2)-receptor antagonist) or NPC567 (another B(2)-receptor...
Serum homocysteine and oxidative stress evaluation during exercise in horse.
Polish journal of veterinary sciences    August 4, 2009   Volume 12, Issue 2 169-174 
Fazio F, Casella S, Giannetto C, Caola G, Piccione G.The aim of the present work was to evaluate the influence of exercise on serum homocysteine and oxidative stress in Thoroughbreds during official races. For our study, ten Thoroughbred mares, 5 years old and clinically healthy, were used. The horses were trained to take part in an official 2100 meter race. Reactive oxygen species (dROMs), antioxidant barrier (Oxy-adsorbent), thiol antioxidant barrier (SHp) and homocysteine (sHcy) were investigated. Blood samples, collected from each horse at rest, immediately after the race, and 30 and 180 min after the end of the race, were collected by jugul...
Electrophysiological characterization of electrolyte and nutrient transport across the small intestine in horses.
Journal of animal physiology and animal nutrition    August 4, 2009   Volume 93, Issue 3 287-294 doi: 10.1111/j.1439-0396.2008.00882.x
Cehak A, Burmester M, Geburek F, Feige K, Breves G.The aim of this study was to characterize the transport mechanisms of electrolytes and nutrients across the jejunum of nine healthy horses electrophysiologically. The stripped mucosa was mounted in Ussing chambers and tissue conductances (G(t)) and short circuit currents (I(sc)) were continuously monitored. After blocking the sodium and potassium channels with amiloride, tetraethylammonium chloride (TEA) and barium, chloride secretion was stimulated by carbachol and forskolin. Subsequently, chloride channels were inhibited by 4,4'-diisothiocyanato-stilbene-2,2'-disulfonic acid, 5-nitro-2-(3-ph...
Influence of age on surfactant isolated from healthy horses maintained on pasture.
Journal of veterinary internal medicine    August 4, 2009   Volume 23, Issue 3 612-618 doi: 10.1111/j.1939-1676.2009.0298.x
Christmann U, Hite RD, Witonsky SG, Elvinger F, Werre SR, Thatcher CD, Tan RH, Buechner-Maxwell VA.Surfactant alterations are described in horses after exercise, anesthesia, and prolonged transport, in horses with recurrent airway obstruction, and in neonatal foals. The effect of horse age or bronchoalveolar lavage fluid (BALF) sample characteristics on surfactant is unknown. Objective: To evaluate surfactant phospholipid composition and function in healthy horses, and to investigate the influence of age and BALF sample characteristics on surfactant. Methods: Seventeen healthy horses 6-25 years of age maintained on pasture year-round. Methods: BALF was collected by standard procedures and w...
Methods and reliability of tissue Doppler imaging for assessment of left ventricular radial wall motion in horses.
Journal of veterinary internal medicine    August 4, 2009   Volume 23, Issue 3 643-652 doi: 10.1111/j.1939-1676.2009.0287.x
Schwarzwald CC, Schober KE, Bonagura JD.Noninvasive assessment of left ventricular (LV) function is incompletely studied in horses. Objective: The goals of this study were to investigate the feasibility, techniques, and reliability of tissue Doppler imaging (TDI) for characterization of LV radial wall motion in healthy horses. Methods: Three Standardbreds, 3 Thoroughbreds; age 8-14 years; body weight 517-606 kg. Methods: Repeated echocardiographic examinations were performed by 2 observers in unsedated horses using TDI. Test reliability was determined by estimating measurement variability, within-day interobserver variability, and b...
Changes in intramuscular amino acid levels in submaximally exercised horses – a pilot study.
Journal of animal physiology and animal nutrition    August 3, 2009   Volume 94, Issue 4 455-464 doi: 10.1111/j.1439-0396.2009.00929.x
van den Hoven R, Bauer A, Hackl S, Zickl M, Spona J, Zentek J.The time-dependent changes in intramuscular amino acid (AA) levels caused by exercise and by feeding a protein/AA supplement were analysed in nine horses. Horses were submitted to a total of four standardized exercise tests (SETs). Amino acid concentrations were determined prior to, immediately after, 4 and 18 h after exercise. The experiment was subdivided into two consecutive periods of 3 weeks. In each period two SETs were performed. In the second period, horses were given a protein/AA supplement within 1 h after exercise. Significant changes in mean plasma AA levels similar to previous stu...
Influence of the position of the foot on MRI signal in the deep digital flexor tendon and collateral ligaments of the distal interphalangeal joint in the standing horse.
Equine veterinary journal    August 1, 2009   Volume 41, Issue 5 498-503 doi: 10.2746/042516409x394472
Spriet M, Zwingenberger A.Hyperintense signal is sometimes observed in ligaments and tendons of the equine foot on standing magnetic resonance examination without associated changes in size and shape. In such cases, the presence of a true lesion or an artifact should be considered. A change in position of a ligament or tendon relative to the magnetic field can induce increased signal intensity due to the magic angle effect. Objective: To assess if positional rotation of the foot in the solar plane could be responsible for artifactual changes in signal intensity in the collateral ligaments of the distal interphalangeal ...
Effects of different forms of exercise on post inhibitory rebound and unwanted behaviour in stabled horses.
Equine veterinary journal    August 1, 2009   Volume 41, Issue 5 487-492 doi: 10.2746/095777309x383883
Freire R, Buckley P, Cooper JJ.It is unknown if different locomotor activities are equally effective at meeting the stabled horse's need for exercise and if they attenuate unwanted behaviour. Objective: Alternative forms of exercise influence the intensity of locomotor activities during a period of turn-out (the so-called rebound effect) and the occurrence of unwanted or undesirable activities during standard handling situations. Methods: Twenty-four horses kept in stables were randomly assigned to one of 4 exercise regimes (walker, treadmill, turn-out and riding) for 4 consecutive days. Because these forms of exercise prov...
Lactobacillus equicursoris sp. nov., isolated from the faeces of a thoroughbred racehorse.
International journal of systematic and evolutionary microbiology    July 31, 2009   Volume 60, Issue Pt 1 109-112 doi: 10.1099/ijs.0.009290-0
Morita H, Shimazu M, Shiono H, Toh H, Nakajima F, Akita H, Takagi M, Takami H, Murakami M, Masaoka T, Tanabe S, Hattori M.We previously isolated five strains of putative lactobacilli from the faeces of a thoroughbred horse (a 4-year-old male). Of the five strains, four were identified as members of existing Lactobacillus species; however, sequence analysis of the 16S rRNA gene revealed that the fifth isolate, DI70(T), showed approximately 97 % identity (1325/1366 bp) with the type strain of Lactobacillus delbrueckii. Therefore, we considered the possibility that DI70(T) represents a novel species of the genus Lactobacillus. Cells of strain DI70(T) were Gram-stain-positive, catalase-negative, non-spore-forming, no...
Reproducibility of a non-invasive ultrasonic technique of tendon force measurement, determined in vitro in equine superficial digital flexor tendons.
Journal of biomechanics    July 31, 2009   Volume 42, Issue 13 2210-2213 doi: 10.1016/j.jbiomech.2009.06.005
Crevier-Denoix N, Ravary-Plumioën B, Evrard D, Pourcelot P.A non-invasive ultrasonic (US) technique of tendon force measurement has been recently developed. It is based on the relationship demonstrated between the speed of sound (SOS) in a tendon and the traction force applied to it. The objectives of the present study were to evaluate the variability of this non-linear relationship among 7 equine superficial digital flexor (SDF) tendons, and the reproducibility of SOS measurements in these tendons over successive loading cycles and tests. Seven SDF tendons were equipped with an US probe (1MHz), secured in contact with the skin overlying the tendon me...
Molecular characterization and expression pattern of the equine lactate dehydrogenase A and B genes.
Gene    July 30, 2009   Volume 447, Issue 1 40-50 doi: 10.1016/j.gene.2009.07.017
Echigoya Y, Sato T, Itou T, Endo H, Sakai T.The species-specific properties of LDH isozymes are essentially determined by M (muscle) and H (heart) subunit proteins encoded by the LDHA and LDHB genes, respectively. In the present study, we molecularly characterized the full-length equine lactate dehydrogenase A (eLDHA) and B (eLDHB) cDNAs. The eLDHA cDNA consisted of a 999-bp open reading frame (ORF), while the eLDHB and newly acquired bat LDHB consisted of a 1002-bp ORF, which is 3 bp shorter than the LDHB ORF of other registered mammals. The alignment of amino acid sequences showed that eLDHA acquired positively charged His 88 and 226,...
Osmotic tolerance limits and membrane permeability characteristics of stallion spermatozoa treated with cholesterol.
Cryobiology    July 29, 2009   Volume 59, Issue 2 201-206 doi: 10.1016/j.cryobiol.2009.07.009
Glazar AI, Mullen SF, Liu J, Benson JD, Critser JK, Squires EL, Graham JK.Stallion spermatozoa exhibit osmotic damage during the cryopreservation process. Recent studies have shown that the addition of cholesterol to spermatozoal membranes increases the cryosurvival of bull, ram and stallion spermatozoa, but the exact mechanism by which added cholesterol improves cryosurvival is not understood. The objectives of this study were to determine if adding cholesterol to stallion sperm membranes alters the osmotic tolerance limits and membrane permeability characteristics of the spermatozoa. In experiment one, stallion spermatozoa were treated with cholesterol-loaded cycl...
The fracture toughness of cancellous bone.
Journal of biomechanics    July 29, 2009   Volume 42, Issue 13 2054-2060 doi: 10.1016/j.jbiomech.2009.06.001
Cook RB, Zioupos P.The mechanical capacity and integrity of cancellous bone is crucial in osteoporosis, a condition which is set to become more prevalent with increasing lifespan and population sizes. The fracture toughness (FT) of cancellous bone has never been examined before and the conditions associated with the growth of a major crack through the lattice of cancellous bone, a cellular solid, may improve our understanding for structural integrity of this material. The aim of this study is to provide (i) basic data on cancellous bone FT and (ii) the experimental support for the hypothesis of Gibson, L.J., Ash...
Comparative study of the characteristics and properties of tendinocytes derived from three tendons in the equine forelimb.
Tissue & cell    July 28, 2009   Volume 42, Issue 1 9-17 doi: 10.1016/j.tice.2009.06.002
Hosaka YZ, Takahashi H, Uratsuji T, Tangkawattana P, Ueda H, Takehana K.The aim of this study was to determine the characteristic differences in tendinocytes derived from tendons in the equine forelimb, superficial digital flexor tendon (SDFT), deep digital flexor tendon (DDFT) and common digital extensor tendon (CDET), in morphology, proliferation, collagen production ability and ability for synthesis of matrix metalloproteinases (MMPs). Significant differences were observed in cell number in vivo. The cellular number was largest in the SDFT and smallest in the CDET. The values of in vitro proliferation ratios and ability for synthesis of collagen and MMPs were l...
Streptomyces atriruber sp. nov. and Streptomyces silaceus sp. nov., two novel species of equine origin.
International journal of systematic and evolutionary microbiology    July 23, 2009   Volume 59, Issue Pt 11 2899-2903 doi: 10.1099/ijs.0.008862-0
Labeda DP, Price NP, Kroppenstedt RM, Donahue JM, Williams NM, Sells SF.Two actinomycete strains, NRRL B-24165(T) and NRRL B-24166(T), isolated from lesions on equine placentas in Kentucky, USA, were analysed using a polyphasic taxonomic approach. On the basis of phylogenetic analysis of 16S rRNA gene sequences, morphological observations and the presence of ll-diaminopimelic acid as the diagnostic diamino acid in whole-cell hydrolysates, the new isolates clearly belonged to the genus Streptomyces. Analyses of the phylogenetic positions of strains NRRL B-24165(T) and NRRL B-24166(T) based on 16S rRNA gene sequences of all recognized species of the genus Streptomyc...
Steroid hormone receptors ERalpha and PR characterised by immunohistochemistry in the mare adrenal gland.
Acta veterinaria Scandinavica    July 22, 2009   Volume 51, Issue 1 31 doi: 10.1186/1751-0147-51-31
Alm YH, Sukjumlong S, Kindahl H, Dalin AM.Sex steroid hormone receptors have been identified in the adrenal gland of rat, sheep and rhesus monkey, indicating a direct effect of sex steroids on adrenal gland function. Methods: In the present study, immunohistochemistry using two different mouse monoclonal antibodies was employed to determine the presence of oestrogen receptor alpha (ERalpha) and progesterone receptor (PR) in the mare adrenal gland. Adrenal glands from intact (n = 5) and ovariectomised (OVX) (n = 5) mares, as well as uterine tissue (n = 9), were collected after euthanasia. Three of the OVX mares were treated with a sing...
Intensity of activation and timing of deactivation modulate elastic energy storage and release in a pennate muscle and account for gait-specific initiation of limb protraction in the horse.
The Journal of experimental biology    July 21, 2009   Volume 212, Issue Pt 15 2454-2463 doi: 10.1242/jeb.027995
Lichtwark GA, Watson JC, Mavrommatis S, Wilson AM.The equine biceps brachii (biceps) initiates rapid limb protraction through a catapult mechanism. Elastic strain energy is slowly stored in an internal tendon and is then rapidly released to protract the forelimb. The muscle fibres are short, have little scope for length change and can therefore only shorten slowly compared with the speed at which the whole muscle must shorten, which makes them poor candidates for driving rapid limb protraction. We suggest that the muscle fibres in the biceps act to modulate the elastic energy output of the muscle-tendon unit (MTU) to meet the demands of locom...
Effects of dermal dexamethasone application on ACTH and both basal and ACTH-stimulated cortisol concentration in normal horses.
Journal of veterinary pharmacology and therapeutics    July 21, 2009   Volume 32, Issue 4 379-387 doi: 10.1111/j.1365-2885.2008.01054.x
Abraham G, Allersmeier M, Gottschalk J, Schusser GF, Hoppen HO, Ungemach FR.There are no data available regarding the systemic (adverse) effects which might be induced by topical/dermal glucocorticoids (GCs) application in the horse. Besides their widespread use for the treatment of a variety of peripheral inflammatory disorders such as atopic dermatitis, eczemas or arthritis in the horse, their surreptitious application has become a concern in doping cases in competition/performance horses. Assessing both basal and ACTH-stimulated plasma cortisol as well as basal ACTH concentrations following application of dexamethsone-containing dermal ointment is necessary to dete...
Modern riding style improves horse racing times.
Science (New York, N.Y.)    July 18, 2009   Volume 325, Issue 5938 289 doi: 10.1126/science.1174605
Pfau T, Spence A, Starke S, Ferrari M, Wilson A.When animals carry loads, there is a proportionate increase in metabolic cost, and in humans this increase in cost is reduced when the load is elastically coupled to the load bearer. Major horse race times and records improved by 5 to 7% around 1900 when jockeys adopted a crouched posture. We show that jockeys move to isolate themselves from the movement of their mount. This would be difficult or impossible with a seated or upright, straight-legged posture. This isolation means that the horse supports the jockey's body weight but does not have to move the jockey through each cyclical stride pa...
A 40-year odyssey into the mysteries of equine luteolysis.
Theriogenology    July 18, 2009   Volume 72, Issue 5 591-598 doi: 10.1016/j.theriogenology.2009.05.016
Ginther OJ.Variation is the principal barrier to progress in unraveling the complexities of biological mechanisms. The resulting slow research progress is well illustrated in the chronology of events in elucidating the mechanism for regression of the corpus luteum (luteolysis) during the equine estrous cycle. Many of the underlying foundations of the female reproductive system in farm animals were developed during the 1930s to 1950s, despite the lack of methods for determining the concentrations of circulating hormones. In the 1960s, a uterine luteolysin was postulated on the basis of several experimenta...
Temporal analysis of equine bone marrow aspirate during establishment of putative mesenchymal progenitor cell populations.
Stem cells and development    July 17, 2009   Volume 19, Issue 2 269-282 doi: 10.1089/scd.2009.0091
Radcliffe CH, Flaminio MJ, Fortier LA.Mesenchymal progenitor cells (MPCs) are often characterized using surface markers after expansion and treatment in culture. There are no studies directly comparing gene and protein markers in undifferentiated samples during the very early phases of culture. The goal of this study was to evaluate temporal gene and protein expression changes during establishment of equine MPC cultures. Bone marrow aspirate was obtained from 35 horses and processed by density gradient centrifugation. In freshly isolated bone marrow, mononuclear cells had variable expression of CD44, CD11a/CD18, CD90, and CD45RB c...
Effect of cryopreservation on nitric oxide production by stallion spermatozoa.
Biology of reproduction    July 15, 2009   Volume 81, Issue 6 1106-1111 doi: 10.1095/biolreprod.109.078220
Ortega Ferrusola C, González Fernández L, Macías García B, Salazar-Sandoval C, Morillo Rodríguez A, Rodríguez Martinez H, Tapia JA, Peña FJ.The ability of stallion spermatozoa to produce nitric oxide (NO) before (fresh) and after freezing and thawing (FT) was evaluated by means of flow cytometry after loading the sperm suspension with the probe, 4,5-diaminofluorescenin diacetate. The presence of NO synthase (NOS) was investigated by Western blotting using anti-NOS1, anti-NOS3, or anti-universal NOS antibodies (Abs). While NO was detected both in fresh and FT sperm suspensions, its production increased after cryopreservation only when egg yolk was removed from the extender. Anti-NOS1 Ab intensively labeled a single band with an app...
Single-layer centrifugation with Androcoll-E can be scaled up to allow large volumes of stallion ejaculate to be processed easily.
Theriogenology    July 14, 2009   Volume 72, Issue 6 879-884 doi: 10.1016/j.theriogenology.2009.05.015
Morrell JM, Johannisson A, Dalin AM, Rodriguez-Martinez H.The objective of the current study was to optimize the volumes of Androcoll-E and sperm sample used in various sizes of centrifuge tube to scale up single-layer centrifugation (SLC) for routine use in the field. Although sperm suspensions of equivalent quality were produced using Androcoll-E in small and large tubes, the sperm yield was much lower in the latter (P<0.001). In contrast, in 200-mL tubes (XL), the yields were approximately 25% higher than those for the small tubes. An increased volume (4.5 mL) of extended ejaculate in small tubes (SLC-Inc) or 15 to 18 mL extended ejaculate on 1...
Preface. New perspectives in equine colic.
The Veterinary clinics of North America. Equine practice    July 8, 2009   Volume 25, Issue 2 xiii-xiv doi: 10.1016/j.cveq.2009.05.002
Andrews FM.No abstract available
Development of a metabonomic approach based on LC-ESI-HRMS measurements for profiling of metabolic changes induced by recombinant equine growth hormone in horse urine.
Analytical and bioanalytical chemistry    July 8, 2009   Volume 394, Issue 8 2119-2128 doi: 10.1007/s00216-009-2912-8
Kieken F, Pinel G, Antignac JP, Monteau F, Christelle Paris A, Popot MA, Bonnaire Y, Le Bizec B.Despite the worldwide existing regulation banning the use of the recombinant equine growth hormone (reGH) as growth promoter, it is suspected to be used in horseracing to improve performances. Various analytical methods previously developed to screen for its misuse have encountered some limitations in terms of detection timeframes, in particular during the first days following reGH administration. A novel strategy involving the characterization of global metabolomic fingerprints in urine samples of non-treated and reGH-treated horses by liquid chromatography-electrospray-high-resolution mass s...
New insights into the mechanisms of fertilization: comparison of the fertilization steps, composition, and structure of the zona pellucida between horses and pigs.
Biology of reproduction    July 8, 2009   Volume 81, Issue 5 856-870 doi: 10.1095/biolreprod.109.077651
Mugnier S, Dell'Aquila ME, Pelaez J, Douet C, Ambruosi B, De Santis T, Lacalandra GM, Lebos C, Sizaret PY, Delaleu B, Monget P, Mermillod P....The mechanism of fertilization remains largely enigmatic in mammals. Most studies exploring the molecular mechanism underlying fertilization have been restricted to a single species, generally the mouse, without a comparative approach. However, the identification of divergences between species could allow us to highlight key components in the mechanism of fertilization. In the pig, in vitro fertilization (IVF) and polyspermy rates are high, and spermatozoa penetrate easily through the zona pellucida (ZP). In contrast, IVF rates are low in the horse, and polyspermy is scarce. Our objective was ...
The new framework for understanding placental mammal evolution.
BioEssays : news and reviews in molecular, cellular and developmental biology    July 8, 2009   Volume 31, Issue 8 853-864 doi: 10.1002/bies.200900053
Asher RJ, Bennett N, Lehmann T.An unprecedented level of confidence has recently crystallized around a new hypothesis of how living placental mammals share a pattern of common descent. The major groups are afrotheres (e.g., aardvarks, elephants), xenarthrans (e.g., anteaters, sloths), laurasiatheres (e.g., horses, shrews), and euarchontoglires (e.g., humans, rodents). Compared with previous hypotheses this tree is remarkably stable; however, some uncertainty persists about the location of the placental root, and (for example) the position of bats within laurasiatheres, of sea cows and aardvarks within afrotheres, and of der...