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Atlas of Clinical Imaging and Anatomy of the Equine Head presents a clear and complete view of the complex anatomy of the equine head using cross-sectional imaging. * Provides a comprehensive comparative atlas to structures of the equine head * Pairs gross anatomy with radiographs, CT, and MRI images * Presents an image-based reference for understanding anatomy and pathology * Covers radiography, computed tomography, and magnetic resonance imaging

Clinical Radiology of the Horse is the best-selling, practical guide to all areas of equine radiography and radiology written by an experienced group of clinicians with a broad range of backgrounds. Offers an atlas of normal and clinical images, as well as a comprehensive guide to techniques, equipment, positioning, and interpretation for general veterinary practitioners and specialists in imaging and orthopaedics Updates to this fourth edition fully reflect the move to digital imaging with many new figures in the book and major revisions to the chapters on the head, thorax, and abdomen Contains expanded coverage of the foot, pastern, and fetlock (now in separate chapters) Includes a password-protected website with all the images from the book as well as over 200 additional images with examples of more subtle lesions, more fractures, correct technique and positioning versus incorrect, immature horses, progression of disease, and pathological images Offers an atlas of normal and clinical images, as well as a comprehensive guide to techniques, equipment, positioning, and interpretation for general veterinary practitioners and specialists in imaging and orthopaedics

The domestication of horses was very important in the history of humankind. However, the ancestry of modern horses and the location and timing of their emergence remain unclear. Gaunitz et al. generated 42 ancient-horse genomes. Their source samples included the Botai archaeological site in Central Asia, considered to include the earliest domesticated horses. Unexpectedly, Botai horses were the ancestors not of modern domestic horses, but rather of modern Przewalski's horses. Thus, in contrast to current thinking on horse domestication, modern horses may have been domesticated in other, more Western, centers of origin. Science , this issue p. 111 The earliest herded horses were ancestors of feral Przewalski’s horses but not of modern domesticated horses. The Eneolithic Botai culture of the Central Asian steppes provides the earliest archaeological evidence for horse husbandry, ~5500 years ago, but the exact nature of early horse domestication remains controversial. We generated 42 ancient-horse genomes, including 20 from Botai. Compared to 46 published ancient- and modern-horse genomes, our data indicate that Przewalski’s horses are the feral descendants of horses herded at Botai and not truly wild horses. All domestic horses dated from ~4000 years ago to present only show ~2.7% of Botai-related ancestry. This indicates that a massive genomic turnover underpins the expansion of the horse stock that gave rise to modern domesticates, which coincides with large-scale human population expansions during the Early Bronze Age.

Significance The domestication of the horse revolutionized warfare, trade, and the exchange of people and ideas. This at least 5,500-y-long process, which ultimately transformed wild horses into the hundreds of breeds living today, is difficult to reconstruct from archeological data and modern genetics alone. We therefore sequenced two complete horse genomes, predating domestication by thousands of years, to characterize the genetic footprint of domestication. These ancient genomes reveal predomestic population structure and a significant fraction of genetic variation shared with the domestic breeds but absent from Przewalski’s horses. We find positive selection on genes involved in various aspects of locomotion, physiology, and cognition. Finally, we show that modern horse genomes contain an excess of deleterious mutations, likely representing the genetic cost of domestication. The domestication of the horse ∼5.5 kya and the emergence of mounted riding, chariotry, and cavalry dramatically transformed human civilization. However, the genetics underlying horse domestication are difficult to reconstruct, given the near extinction of wild horses. We therefore sequenced two ancient horse genomes from Taymyr, Russia (at 7.4- and 24.3-fold coverage), both predating the earliest archeological evidence of domestication. We compared these genomes with genomes of domesticated horses and the wild Przewalski’s horse and found genetic structure within Eurasia in the Late Pleistocene, with the ancient population contributing significantly to the genetic variation of domesticated breeds. We furthermore identified a conservative set of 125 potential domestication targets using four complementary scans for genes that have undergone positive selection. One group of genes is involved in muscular and limb development, articular junctions, and the cardiac system, and may represent physiological adaptations to human utilization. A second group consists of genes with cognitive functions, including social behavior, learning capabilities, fear response, and agreeableness, which may have been key for taming horses. We also found that domestication is associated with inbreeding and an excess of deleterious mutations. This genetic load is in line with the “cost of domestication” hypothesis also reported for rice, tomatoes, and dogs, and it is generally attributed to the relaxation of purifying selection resulting from the strong demographic bottlenecks accompanying domestication. Our work demonstrates the power of ancient genomes to reconstruct the complex genetic changes that transformed wild animals into their domesticated forms, and the population context in which this process took place.

A reduction in the number of digits has evolved many times in tetrapods, particularly in cursorial mammals that travel over deserts and plains, yet the underlying developmental mechanisms have remained elusive. Here we show that digit loss can occur both during early limb patterning and at later post-patterning stages of chondrogenesis. In the ‘odd-toed’ jerboa ( Dipus sagitta ) and horse and the ‘even-toed’ camel, extensive cell death sculpts the tissue around the remaining toes. In contrast, digit loss in the pig is orchestrated by earlier limb patterning mechanisms including downregulation of Ptch1 expression but no increase in cell death. Together these data demonstrate remarkable plasticity in the mechanisms of vertebrate limb evolution and shed light on the complexity of morphological convergence, particularly within the artiodactyl lineage. A study of limb development in multiple mammals reveals that evolutionary digit loss has occured in many different ways—at different stages and by different mechanisms, such as regulation of Shh in initial digit specification events or by removal of digits through cell death. Mechanisms of evolutionary limb loss The basic five-digit limb of tetrapods has been altered many times and in many ways during evolution, usually by the progressive loss of digits. Two papers published in this issue of Nature examine the developmental changes underlying digit reduction in mammals. Javier Lopez-Rios et al . look at cattle, where digits three and four are modified to form hooves; digits two and five are vestigial, and the first digit is lost. The first limb bud is shown to be progressively lost as it develops. The Ptch1 gene, which encodes a receptor for the limb-development morphogen Sonic hedgehog (SHH), is upregulated due to evolutionary alteration of a Ptch1 cis -regulatory module that no longer responds to graded SHH signalling during bovine handplate development. Kimberly Cooper et al . show, using a wide range of mammals, that mechanisms of digit loss vary in different lineages. In creatures as varied as the jerboa and the camel, cell death sculpts the tissue in the emerging limb to leave the remaining toes. In other creatures, such as the pig, digit loss is orchestrated by earlier limb patterning events with no increase in cell death. Taken together these findings demonstrate remarkable plasticity in the mechanisms of limb evolution in hooved mammals and rodents, yet reveal a degree of evolutionary convergence.

The evolution of high-crowned molars among horses (Family Equidae) is thought to be an adaptation for abrasive diets associated with the spread of grasslands. The sharpness and relief of the worn cusp apices of teeth (mesowear) are a measure of dietary abrasion. We collected mesowear data for North American Equidae for the past 55.5 million years to test the association of molar height and dietary abrasion. Mesowear trends in horses are reflective of global cooling and associated vegetation changes. There is a strong correlation between mesowear and crown height in horses; however, most horse paleopopulations had highly variable amounts of dietary abrasion, suggesting that selective pressures for crown height may have been weak much of the time. However, instances of higher abrasion were observed in some paleopopulations, suggesting intervals of stronger selection for the evolution of dentitions, including the early Miocene shortly before the first appearance of Equinae, the horse subfamily in which high-crowned dentitions evolved.