1. Introduction
Adequate sole depth in the hooves of domesticated, barefoot-kept horses is essential for locomotion. In healthy equine feet, the sole cushions protect the distal phalanx [
1] and are typically thicker caudally [
2]. Although there is no standardized sole depth for individual barefoot-kept breeds, conflicting information exists regarding the optimal internal thickness, ranging between 10˗15 mm [
3,
4,
5]. Nonetheless, Mellish et al. [
6] conducted a study on cadaver hooves from feral horses in Canada and observed shallow soles, with 65% having less than 11 mm at the toe. Sole depth at the palmar process (PP) was similar, revealing negative palmar angles. These herds resided on soft substrate, yet remained sound, indicating sole adaptation [
6]. Feral horses in New Zealand presented similar results, displaying hoof morphology adapted to their soft environments [
7]. Radiographic imaging of the soles of live animals, including both the vascular dermis and the thicker keratinized epidermis, does not consider possible preferential contributions of these tissue elements to foot health and soundness.
Understanding the mechanical and biological properties of the equine foot is crucial for comprehending sole function and growth. Trimming methods that distribute impact forces between the hoof wall, sole, and frog to the appropriate internal foot tissues, are essential for achieving efficient and correct biomechanics for locomotion [
4,
8]. Horses with some degree of long toes and varying underrun heels, while considered sound, may not have healthy feet [
9]. This hoof shape is typical of navicular cases [
10]. Although the hoof can twist, torque, flex, and expand under varying impact and loading forces over uneven ground surfaces [
8,
11], its capability to correctly process these energy waves minimizes foot pathologies. The somatosensory system detects and responds to such forces, becoming an important contributor to sole responses, including growth. Foot receptors were mapped within the hoof 30 years ago, illustrating their potential neurophysiological roles in movement [
12]. However, their potential involvement in other solar actions, particularly in guiding and modulating energy waveforms through various foot tissues, vascular perfusion, and even foot growth on hard surfaces, has never been explored. This neuroanatomical system, rich in receptors allowing the foot to obtain information from the terrain on which it moves [
13], senses specific sensations, including pressure, vibration, balance, pain, and temperature, all of which contribute directly and indirectly to the overall internal health of the foot [
13]. Particular fast-adapting receptors known as Pacinian corpuscles (or FA2 receptors), mainly located caudally, respond to pressure and vibration, contributing to increased perfusion of matrix tissues, to enhance sole growth [
13]. In habitually barefoot human populations, thicker soles are evident at the footpad [
14], owing to a large cluster of FA2s at this site [
15].
Diet and management are critical factors affecting sole health. A high sugar-starch diet based on grains and concentrated feed, prolonged periods of grazing on lush grass, and obesity, can cause metabolic issues, leading to laminitis or founder [
16,
17]. Very thin soles are often associated with this pathology, which can eventually immobilize the horses [
9,
17]. Unhygienic housing conditions that confine horses to standing in their urine and manure for extended periods or muddy environments can also lead to sole tenderness and thrush [
4,
18].
Internal sole measurements, including the positions of the phalanges, palmar and plantar angles of the distal phalanx, and mediolateral imbalance [
19], can be confirmed through radiographs [
3,
4,
20].
Recent studies emphasize the health benefits of unshod horse hooves [
21,
22,
23,
24], with an increasing number of horse owners choosing barefoot options [
25,
26]. To the best of our knowledge, sole thickness in hooves of domesticated, barefoot-kept horses walking over hard, artificial environments has not been reported in equine veterinary literature. Although nineteenth-century literature offers insights into the capabilities of hooves of barefoot-kept horses ridden or driven over dirt and cobbled roads [
27,
28], prior to the introduction of tarmac [
29], none of these studies measured sole depth.
In this study, we aimed to use completely forage-fed barefoot-kept equines with a 24/7 turnout and radiographically measure the sole depth, including the external morphological characteristics of their hooves, in response to three abrasive surfaces (30 km tarmac, 8 km concrete screed, and 12 km limestone gravel). As only cadaveric specimens have been used to assess the morphological characteristics of this solar area to date, we hypothesized that the soles of the hooves would become hard (calloused), would require trimming, and that sole depth would decrease slightly by the end of the study.
4. Discussion
The outcomes of the present study were counterintuitive. Despite the development of hardened soles, the unexpected findings included hoof wall trimming on day 9, phalangeal misalignment, and sole growth. In the radiographs, sole thickness was more noticeable at the PP in eleven hooves, yet external measurements presented an increase in heel height in eight feet of both horses and ponies. Bellenzani et al. [
11] observed barefoot horses exercising on a treadmill for four weeks at different gaits, reporting higher heels; however, they did not explain the observed PP growth. In our study, the horses exhibited a more pronounced, natural concavity in the front hooves, confirming a thicker sole depth at the apex, evident in radiographs. Most external measurements from the coronet-toe were shorter in both horses and ponies, suggesting wear at the toes during the breakover phase, on abrasive surfaces. Frog apex-toe measurements indicated decreased distances in all equines, signifying wear at the toes (Figure 8).
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Accelerated hoof wall growth became evident in the outer hoof capsules after covering 36 km. An hour of walking per day on abrasive surfaces may not sufficiently wear away the hoof wall. However, our results emphasize the importance of short trimming cycles of no longer than three weeks. Follow-up radiographs revealed misaligned phalanges in four hooves after the 50-km walk. These animals are maintained on a regular 2˗3-week trimming cycle to minimize hoof wall loading and sustain balance. Although literature highlights the importance of short trimming cycles in horses, especially those with hoof pathology [
4], practices in the UK often utilize 6˗8 week trimming cycles, for both shod and barefoot horses [
31]; however, there is no information on phalangeal misalignment. Kummer et al. [
32] reported regular, shoeing cycles of 8˗10 weeks but did not provide any information on palmar/plantar angles or mediolateral imbalance. Further studies are required to explore misalignment and mediolateral imbalances during extended trimming cycles. While our findings highlight the benefits of shorter trimming cycles, hoof wall growth must be examined in a larger population of barefoot-kept horses. To date, the effects of climate, housing conditions, diet, and exercise surface on hoof wall growth has not been documented.
A comprehensive investigation was essential to investigate accelerated sole growth, given the limited information in equine veterinary literature, particularly owing to the constraints of conventional radiography in visualising soft tissue. Scientific studies indicate that the hoof serves as a neurosensory organ, rich in sensory nerves with various fiber types and neurotransmitters, transmitting information to the central nervous system [
13]. These neurotransmitters act as receptors processing environmental stimuli [
13], mirroring functions observed in the human foot [
33]. Microscopic examination of internal hoof anatomy reveals Pacinian corpuscles (mechanoreceptors) abundant in the caudal part of the foot, potentially influencing vascular changes [
34]. They require constantly changing stimuli, respond to pressure and vibration, and are crucial in the gait cycle of a horse. Since these corpuscles are scant in the toe region or corium, this indicates that the horse is a rear-foot striker [
13]. However, the effects of these corpuscles on sole depth in horses walking over hard surfaces have not been investigated. In humans, Pacinian corpuscles are tightly grouped in various soft tissues around the metatarsal pad and proximal phalanges [
15]. These FA2s are higher in number than other receptor-types located in the foot [
14] and may suggest that humans are foot-first strikers when running and jumping, landing first on their foot pads to minimize impact and dampen the ground reaction forces (GRFs) [
35]. Sole growth was more prominent at the PP of the animals used in the study.
Given that most Pacinian corpuscles are located at the caudal part of the hoof, respond to high-frequency vibrations and rapidly adapt to pressure and stimuli [
13], growth in this area was accelerated as it constituted a primary impact point on hard surfaces. A study conducted on individuals in Western Kenya who were habitually barefoot, revealed thicker calluses at the metatarsal head than at the heel [
14], suggesting a parallel function of FA2s at the site, upon impact. Here, the solar side of the foot produces calluses through environmental stimuli, simultaneously protecting against injury [
35].
Conversely, when assessing sole depth in the hooves of Sable Island and Kaimanawa feral horses, they appeared thin, both at the crena and PP, irrespective of overgrown hoof walls. Pacinian corpuscles in this scenario, might perceive or detect less pressure and vibration owing to the tendency of the soft ground to muffle, or mute, impacted vibrations. Furthermore, chronic exposure to soft surfaces could render FA2s less responsive or cause them to atrophy, both of which could lead to reduced afferent activity from the GRF. Additional studies are imperative to explore this phenomenon of thicker sole depths on harder surfaces, independent of hoof wall condition. The potential role of the somatosensory system in modulating changes in the equine foot on different terrains necessitates further investigation.
Ventral to the distal phalanx, sole thickness, as seen radiographically, consists of an internal and external zone of the dermis and keratinized epidermis, respectively. The dermal zone is composed of a microvasculature system, matrix tissues, including elastic and reticular fibers, fascial sheets, and cords along with isolated small clusters of adipocytes. The external layer, composed of epidermal tubules and the intertubular horn, closely adheres to this dermal configuration. Collectively, these elements gradually arch across the width of the foot, between the toe and dorsal third of the frog, whereas caudally, a greater arch of the sole slants toward the epidermal bars juxtaposed near the midline. This complex arrangement, functions as a unique unit, supporting energy dissipation, regenerative capabilities of the solar foot, and directing energy impact forces toward appropriate tissues, including limb loading of the bony column. Within the dermis, a dense microvasculature concentrated at its solar margins and caudally ventral to the palmar processes [
36], serves as a loading site along the peripheral sole, simultaneously contributing to these functions. These two solar layers, either together or alone, may have contributed to the increased sole thickness in horses and ponies walking on the road. This distinctive structural arrangement of solar tissues between the distal phalanx and varied terrain features, highlights the need for further investigation in both of these areas.
The digital cushion (DC) is a wedge-shaped fibrous and fibrocartilage tissue located proximal to the frog, encompassing much of the caudal part of the foot. It plays a critical role in the hoof, buffering high-impact forces in the limbs [
37,
38]. This particular structure absorbs GRFs during heel-first landings, dissipates energy and rebounds back into its shape as the heel is lifted [
38]. Notably, the DC harbors the most neuroreceptors in the foot [
13]. When the frog makes contact with the ground, the DC can receive adequate stimuli, encouraging the development of fibrocartilage [
38,
39]. Thus, the DC may have also contributed to the observed sole thickness in the hooves of the animals in this study. Further investigations could measure digital cushion thickness to confirm this, employing digital sonography, as previously conducted on bovines [
40]. Additionally, considering that the forelimbs of equines support 60% of their body weight, it is plausible that they would possess thicker digital cushions than the hind feet, as documented in bovines [
40].
Load distribution within the equine feet holds paramount importance for locomotion. Trimming techniques should ensure or enhance an even distribution of load among the hoof wall, sole, frog, and bars [
4], with a focus on maintaining short toes to maximize breakover and facilitate heel-first landings. In a well-balanced foot, internal and external structures synergize, fostering compliance, over uneven surfaces. A study on elephants demonstrated that the DC aids in distributing load evenly within the sole, effectively buffering impact forces [
41]. Similar emphasis on the significance and role of a healthy DC has been noted in studies on confined dairy cows [
40,
42,
43]. Nonetheless, we acknowledge that horses did not evolve to roam over artificial surfaces. Instead, their evolution is tailored to terrains, allowing dirt (earth) packs underneath their hooves, facilitating central limb loading and even energy dissipation [
44,
45].
Internally, a well-balanced hoof aligns all three phalanges [
32], with palmar or plantar angles varying between 3-5 degrees [
3]. In follow-up radiographs of the horses in this study, internal sole thickness at the crena ranged between 11-15 mm (mean: 13 mm) in the forefeet with no signs of lameness or sensitivity, especially on gravel. A similar observation applied to ponies, exhibiting a slightly thicker sole, ranging between 14˗16 mm (mean: 15 mm) in the forefeet, at the same anatomical site. Despite the ponies covering more paces owing to their shorter stride throughout the 50 km distance, they displayed thicker soles at the crena. Thick soles, as evidenced in this study, are associated with healthy hooves [
6]. However, broader studies on a larger cohort of barefoot-kept horses are essential to establish a standard optimal sole depth for various surface types. In a well-developed foot, the lateral cartilages should constitute no less than one-third of the total length from the crena to the heel [
4,
44], indicating that a larger area at the caudal part of the foot as observed in radiographs, the healthier. This is evident in the feet of all equines used in the study, hence promoting sole growth. Figure 9 illustrates a radiograph of the Standardbred’s foot.
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Nutrition significantly influences the health and welfare of horses, including their soles [
4,
45,
46]. Additionally, a high sugar-starch diet has long-term implications for hoof health [
9,
47,
48]. Podiatry studies over the last two decades, emphasize the importance of a forage-based diet for optimal hoof health. However, references to this diet predominantly pertain to rehabilitation cases, given that hoof issues often have metabolic origins [
4,
16]. Anecdotal information suggests that many barefoot-kept horses without hoof problems are transitioning to 100% forage-based diets, consisting of good quality hay (tested for nutrients) and replacing grain or pelleted feed with forage balancers [
31], based on the National Research Council recommendations [
49]. These balancers, supplied in powdered form, contain minerals (and some vitamins) typically deficient in hay [
50]. A forage-based diet could be a contributing factor to the intact and distortion-free hooves observed in the animals from this study.
Effective management plays a crucial role in sustaining healthy soles. Barefoot-kept horses benefit from exposure to both hard and soft ground. The more varied the terrain, the more conditioned and resilient their hooves become. The equines included in this study were predominantly outdoors, mainly on compacted soil and grassland (rocky steppe) with minimal green grass. Despite covering a distance of 50 km on hard ground, this exercise may prove less advantageous for their limbs in the long term, especially for horses with arthritis (see Figures 3, 4 & 7) given the heightened concussion absorbed during impact on firm surfaces [
51,
52]. The GRF might predispose them to soft tissue injuries and degenerative joint disease [
52], unless their feet are shielded by dampening products such as hoof boots. Conversely, brief periods of road walking could offer benefits during the shod-to-barefoot transition phase, potentially accelerating the process to increase sole thickness, as demonstrated in this study.
While the horses and ponies in this investigation were hand-walked over abrasive surfaces, we hypothesized that hoof morphology would exhibit minimal variation even if they were ridden. Feral herds traversing several kilometers over challenging terrains, including pregnant mares [
53] that undergo a 15% increase in body weight during the final trimester [
54], with foals weighing around 52 kg at birth [
55], do not experience physical handicaps owing to excess weight, and their hooves remain unaffected.