Horses communicate with eyes and mobile ears

Author Jennifer Wathan is shown with study participant Bartie. Credit: Jennifer Wathan

Horses are sensitive to the facial expressions and attention of other horses, including the direction of the eyes and ears. The findings, reported in the Cell Press journal Current Biology on August 4, are a reminder for us humans to look beyond our own limitations and recognize that other species may communicate in ways that we can't, the researchers say. After all, human ears aren't mobile.

"Our study is the first to examine a potential cue to attention that humans do not have: the ears," says Jennifer Wathan of the University of Sussex. "Previous work investigating communication of attention in animals has focused on cues that humans use: body orientation, head orientation, and eye gaze; no one else had gone beyond that. However, we found that in horses their ear position was also a crucial visual signal that other horses respond to. In fact, horses need to see the detailed facial features of both eyes and ears before they use another horse's head direction to guide them."

The new study also challenges the earlier held notion that animals with eyes to the sides of their heads cannot glean information based on the direction of one another's gaze.

Wathan and the study's senior author Karen McComb took photographs to document cues given by horses when they were paying attention to something. Then Wathan and McComb used those photographs as life-sized models for other horses to look at as they chose between two feeding buckets. In each case, the horse in the photo was paying attention to one of the buckets and not the other. In some instances, the researchers also manipulated the image to remove information from key facial areas, including the eyes and the ears.

The researchers' observations show that horses rely on the head orientation of their peers to locate food. However, that ability to read each other's interest level is disrupted when parts of the face -- the eyes and ears -- are covered up with masks. The ability to correctly judge attention also varied depending on the identity of the horse pictured, suggesting that individual facial features may be important, the researchers report.

Wathan and McComb plan to continue to explore facial features related to the expression of emotion in their horses, noting that horses' rich social lives and close relationship to humans make them particularly interesting as study subjects. Our understanding of horses' social lives might also have implications for their welfare.

"Horses display some of the same complex and fluid social organization that we have as humans and that we also see in chimpanzees, elephants, and dolphins," Wathan says. "The challenges that living in these societies create, such as maintaining valuable social relationships on the basis of unpredictable interactions, are thought to have promoted the evolution of advanced social and communicative skills. There is a general interest in studying species with this social structure."

Source: Cell Press

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Do women and men ride differently? Horses cannot tell the difference

Horses seem to be truly gender-neutral. It doesn't matter to them if their human partner is female or male.
Credit: Juliane Kuhl / Vetmeduni Vienna

Scientists at the Vetmeduni in Vienna have analysed how horses are affected by the sex of their riders. Various parameters of stress were determined in horses and their riders when they covered an obstacle course. The results were surprising: the level of stress on a horse is independent of whether a man or a woman is in the saddle. Furthermore, the stress responses of male and female riders are essentially the same. The results have been published in the Journal of Comparative Exercise Physiology.

For centuries, horse riding was largely restricted to males. The previous situation is in stark contrast to the present day, when nearly 80 percent of riders are women. Modern-day equestrian sports are unique in that men and women compete directly against one another at all levels, from beginners in gymkhanas to national champions in the Olympic Games. "For this reason it is interesting to consider whether a theory of riding that was developed exclusively for men can be applied to women," explains Natascha Ille, the first author of the recent publication.

A rider is a rider

As Ille notes, "It is often assumed that women are more sensitive towards their horses than men. If this is so, male and female riders should elicit different types of response from their horses." Ille, Christine Aurich and colleagues from the Vetmeduni Vienna´s Graf Lehndorff Institute tested this notion by examining eight horses and sixteen riders, including eight men and eight women. Each horse had to jump a standard course of obstacles twice, ridden once by a male and once by a female of similar equestrian experience. The scientists monitored the levels of stress in the horses and their riders, checking the amounts of cortisol in the saliva and the heart rates.

The results were unexpected. The level of cortisol in horses' saliva increased during the test but the increase was not affected by the sex of the rider. The horses' heart rates also increased as a result of taking the course but the increase was irrespective of the human partner in the saddle. The tests on the riders gave similar conclusions. Again, the level of cortisol in the saliva increased but there was no difference between men and women. The riders' pulses sped up when the horses switched from a walk to a canter and accelerated further during the jumping course. But the heart rate curves for male and female riders were close to identical.

The distribution of saddle pressure is the same for male and female riders
In a second experiment, Ille and her colleagues studied the pressure exerted on a horse's back via the saddle. As she explains, "Depending on the rider's posture and position, the pattern of pressure on the horse's back may change dramatically." A special pad placed directly under the saddle was used to analyse saddle pressure in walk, trot and canter. Because female riders are generally lighter than males, the saddle pressure was lower when horses were ridden by females. However, the distribution of pressure did not differ and there was no evidence of differences in the riding posture between males and females.

A fair chance for all

So what does all this mean for modern equestrian sports? Aurich is keen to reassure potential competitors that horses are truly gender-neutral. As she puts it, "Assuming that there is no difference in riding ability, from the horse's point of view, it does not seem to matter whether the human partner is male or female. Our results make it extremely unlikely that horses have a preference for riders of one sex over the other. And when male and female riders compete against one another in equestrian sports, all of them have similar chances of doing well."

Source: Veterinärmedizinische Universität Wien

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Reshaping the horse through millennia: Sequencing reveals genes selected by humans in domestication

A man catches domestic Mongolian horses with a lasso in Khomiin Tal, Mongolia. Credit: Copyright: Ludovic Orlando.

Whole genome sequencing of modern and ancient horses unveils the genes that have been selected by humans in the process of domestication through the latest 5,500 years, but also reveals the cost of this domestication. A new study led by the Centre for GeoGenetics at the University of Copenhagen, in collaboration with scientists from 11 international universities, reports that a significant part of the genetic variation in modern domesticated horses could be attributed to interbreeding with the descendants of a now extinct population of wild horses. This population was distinct from the only surviving wild horse population, that of the Przewalski's horses. The study has been published in the scientific journal Proceedings of the National Academy of Sciences (PNAS).

The domestication of the horse some 5,500 years ago ultimately revolutionized human civilization and societies. Horses facilitated transportation as well as the circulation of ideas, languages and religions. Horses also revolutionized warfare with the advent of chariotry and mounted cavalry and beyond the battlefield horses greatly stimulated agriculture. However, the domestication of the horse and the subsequent encroachment of human civilization also resulted in the near extinction of wild horses.

The only surviving wild horse population, the Przewalski's horses from Mongolia, descends from mere 13 individuals, preserved only through a massive conservation effort. As a consequence of this massive loss of genetic diversity, the effects of horse domestication through times have been difficult to unravel on a molecular level. Says Dr. Ludovic Orlando, Associate Professor at the Centre for GeoGenetics, who led this work

"The classical way to evaluate the evolutionary impact of domestication consists of comparing the genetic information present amongst wild animals and their living domesticates. This approach is ill suited to horses as the only surviving population of wild horses has experienced a massive demographic decline in the 20th century. We therefore decided to sequence the genome of ancient horses that lived prior to domestication to directly assess how pre-domesticated horses looked like genetically."

Recent advances in ancient DNA research have opened the door for reconstructing the genomes of ancient individuals. In 2013, Ludovic Orlando and his team succeeded in decoding the genome of a ~700,000 year-old horse, which represents the oldest genome sequenced to date. This time, the researchers focused on much more recent horse specimens, dating from ~16,000 and ~43,000 years ago. These were carefully selected to unambiguously predate the beginning of domestication, some 5,500 years ago. The bone fossils were excavated in the Taymyr Peninsula, Russia, where arctic conditions favor the preservation of DNA.

The human reshaping of the horse

While the horse contributed to reshaping human civilization, humans in turn reshaped the horse to fit their diverse needs and the diverse environments they lived in. This transformation left specific signatures in the genomes of modern horses, which the ancient genomes helped reveal. The scientists were able to detect a set of 125 candidate genes involved in a wide range of physical and behavioral traits, by comparing the genomes of the two ancient horses with those of the Przewalski's horse and five breeds of domesticated horses. Says Dr. Dan Chang, post-doctoral researcher at the UCSC Paleogenomics Lab and co-leading author of the study:

"Our selection scans identified genes that were already known to evolve under strong selection in horses. This provided a nice validation of our approach."

Dr. Beth Shapiro, head of the UCSC Paleogenomics Lab continues: "We provide the most extensive list of gene candidates that have been favored by humans following the domestication of horses. This list is fascinating as it includes a number of genes involved in the development of muscle and bones. This probably reveals the genes that helped utilizing horses for transportation."

And Dr. Ludovic Orlando from the Centre for GeoGenetics at the University of Copenhagen concludes: "Perhaps even more exciting as it represents the hallmark of animal domestication, we identify genes controlling animal behavior and the response to fear. These genes could have been the key for turning wild animals into more docile domesticated forms."

The 'cost of domestication' in horses

However, the reshaping of the horse genome during their domestication also had significant negative impacts. This was apparent in the increasing levels of inbreeding found amongst domesticates, but also through an enhanced accumulation of deleterious mutations in their genomes relative to the ancient wild horses. This finding supports an earlier theory coined 'the cost of domestication', which predicted increasing genetic loads in domesticates compared to their wild ancestors. Says Professor Laurent Excoffier, University of Bern and group leader at the Swiss Institute for Bioinformatics:

"Domestication is generally associated with repeated demographic crashes. Yet, mutations that negatively impact genes are not eliminated by selection and can even increase in frequency when populations are small. Domestication thus generally comes at a cost, as deleterious mutations can accumulate in the genome. This had already been shown for rice and dogs. Horses now provide another example of this phenomenon."

This is something that was only detectable in the horse in comparison to the ancient genomes, as Przewalski's horses were found to show a proportion of deleterious mutations similar to domesticated horses. Says Hákon Jónsson, PhD-student at the Centre for GeoGenetics, co-leading author of the study: "The recent near extinction of the Przewalski's horse population resulted in the persistence of deleterious mutations in the population, following the same mechanism that once led to the accumulation of deleterious mutations in the genomes of domesticated horses. What is striking is that a similar order of magnitude 
was reached even though this occurred in a much shorter time scale than domestication."

An ancient contribution to the present

In addition, comparison of the ancient and modern genomes revealed that the ancient individuals contributed a significant amount of genetic variation to the modern population of domesticated horses, but not to the Przewalski's horses. This suggests that restocking from a wild population descendant from the ancient horses occurred during the domestication processes that ultimately led to the modern domesticated horses. Mikkel Schubert, PhD- student at the Centre for GeoGenetics, co-leading author of the study concludes:

"This confirms previous findings that wild horses were used to restock the population of domesticated horses during the domestication process. However, as we sequenced whole genomes, we can estimate how much of the modern horse genome has been contributed through this process. Our estimate suggests that at least 13%, and potentially up to as much as 60%, of the modern horse genome has been acquired by restocking from the extinct wild population. That we identified the population that contributed to this process demonstrates that it is possible to identify the ancestral genetic sources that ultimately gave rise to our domesticated horses."

Source: Faculty of Science - University of Copenhagen

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Birth of four foals from genotyped, cryopreserved embryos: A first in Europe

Genotyping allows scientists to choose the embryos they want to use based on different criteria: sex, like in this experiment, the absence of known genetic disorders, or, perhaps in the future, other traits that are tied to behavior, such as emotivity or sociability. Credit: INRA/C. Maitre

IFCE and INRA announced that, for the first time ever in Europe, four foals were successfully born as the result of the transfer of genotyped and cryopreserved embryos. The goal of this work is to better understand embryonic development, control livestock reproduction, and maintain breed genetic diversity. Furthermore, it is advantageous for the horse industry to be able to determine the traits of a future foal.

The INRA Val de Loire center at Nouzilly is where the technology to maintain embryo viability following genotyping and cryopreservation was honed, and then, last summer, the transfer of several embryos took place at the IFCE Haras du Pin Stud Farm, located in the French department of Orne. The partnership between the two institutions has now been cemented by the birth of the healthy foals.

What were the steps leading up to the birth of these foals?

Seven days after fertilization, embryos were collected from Welsh B ponies that are part of INRA's livestock. The embryos were genotyped: scientists sampled some of the embryos' cells to analyze their genomes. In this experiment, embryos were selected based on sex, the idea being to use sex-based selection to test the technique's feasibility. The embryos were then cryopreserved in liquid nitrogen (at -196°). Last summer, they were transferred into saddlebred mares at the Haras du Pin center. After an 11-month gestation period, the foals were born in May. They were of the expected sex: two females and two males. This is the first time that such an event has taken place in Europe, and it is the product of more than 10 years of various types of embryonic research carried out by INRA and IFCE scientists.

What made it so difficult to apply this technique to horses?

Although embryo preservation techniques are already well developed for bovines, small ruminant species, and even humans, preserving horse embryos is a very complex process. For instance, horse embryos vary greatly in size: 7-day-old embryos range in diameter from 200 to 700 micrometers. It is very difficult to cryopreserve the largest embryos because the liquid inside them forms ice crystals when the embryos are frozen at very cold temperatures. What's more, horse embryos are surrounded by a capsule that interferes with successful cryopreservation.

What is the significance of this event, which is the first of its kind in Europe?

There are several reasons why being able to successfully cryopreserve embryos is important. For example, it allows us to maintain breed genetic diversity, particularly that of breeds with small population sizes, such as the Landais or the Poitevin Mulassier. Furthermore, the factor that currently limits the use of embryo transfer is its cost: the transfer center has to maintain a team of recipient mares that are reproductively synchronized with the donor mares. Cryopreservation means that the transfer doesn't have to take place immediately; it can wait until a recipient mare becomes available to receive the embryo. Finally, it may now be possible to directly repopulate horse herds that have experienced losses as a result of various issues, such as disease-related problems, instead of having to use the indirect technique of crossbreeding.

Why genotype the embryos?

Genotyping allows us to choose the embryos we want to use based on different criteria: sex, like in this experiment, the absence of known genetic disorders, or, perhaps in the future, other traits that are tied to behavior, such as emotivity or sociability. It is advantageous for the horse industry to be able to determine the traits of a future foal. We will next aim to simplify the process -- to make this technology more accessible and user friendly for those in the horse industry.

Source: INRA-France

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