Mechanical nociceptive thresholds of dorsal laminae in horses after local anaesthesia of the palmar digital nerves or dorsal branches of the digital nerve.

We examined the hypothesis that the palmar digital nerves (PDNs), but not the dorsal branches (DBs) of the digital nerves, innervate the sensitive dorsal laminae of the equine foot by evaluating the effects of perineural anaesthesia of the PDNs and DBs separately on pain sensation evoked via mechanical stimulation of the dorsal laminae and other regions of the equine foot. Six clinically normal mares were used in a crossover design. A portable dynamometer was used to evaluate mechanical nociceptive thresholds at different points on the dorsal laminae, bulbs of the heel, coronary band and sole before and after the horses underwent perineural injection of PDNs or DBs with a local anaesthetic solution (treated group) or an isotonic saline solution (control group). Cornified tissue was removed from the sole and the dorsal aspect of the hoof wall before evaluations of mechanical nociceptive thresholds. Anaesthetising PDNs distal to the DBs increased mechanical nociceptive thresholds compared to baseline values (P <0.001) at sites assessed in the dorsal laminae, sole, and the bulbs of the heels. Anaesthetising DBs increased mechanical nociceptive thresholds compared to baseline values (P <0.01) only at sites assessed at the most proximal aspect of the foot (i.e., coronary band sites). In conclusion, PDNs, not DBs, are primarily responsible for pain signal transmission evoked by pressure in the dorsal laminae of the foot of clinically normal horses.

Hypergravity within a critical period impacts on the maturation of somatosensory cortical maps and their potential for use-dependent plasticity in the adult.

We investigated experience-dependent plasticity of somatosensory maps in rat S1 cortex during early development. We analyzed both short- and long-term effects of exposure to 2G hypergravity (HG) during the first 3 postnatal weeks on forepaw representations. We also examined the potential of adult somatosensory maps for experience-dependent plasticity after early HG rearing. At postnatal day 22, HG was found to induce an enlargement of cortical zones driven by nail displacements and a contraction of skin sectors of the forepaw map. In these remaining zones serving the skin, neurons displayed expanded glabrous skin receptive fields (RFs). HG also induced a bias in the directional sensitivity of neuronal responses to nail displacement. HG-induced map changes were still found after 16 wk of housing in normogravity (NG). However, the glabrous skin RFs recorded in HG rats decreased to values similar to that of NG rats, as early as the end of the first week of housing in NG. Moreover, the expansion of the glabrous skin area and decrease in RF size normally induced in adults by an enriched environment (EE) did not occur in the HG rats, even after 16 wk of EE housing in NG. Our findings reveal that early postnatal experience critically and durably shapes S1 forepaw maps and limits their potential to be modified by novel experience in adulthood.

The Macroanatomy of the Sacral Plexus and Its Nerves in Eurasian Eagle Owls (Bubo bubo).

This study was carried out to reveal the formation of the sacral plexus in the Eurasian Eagle Owls (Bubo bubo) and the nerves originating from this plexus. Five EEOs, three of them were male and two were female, were provided from Wildlife Rescue and Rehabilitation Center of Kafkas University and used as materials. Following the euthanizing of the animals, abdominal cavity was opened. The nerves of plexus sacrales were dissected and photographed. It was detected that the sacral plexus was formed by the ventral ramus of five synsacral nerves. Moreover, it was determined that the roots of the sacral plexus formed three trunks: the truncus cranialis, the truncus medius and the truncus caudalis in fossa renalis. The availability of the n. ischiofemoralis and the availability of n. parafibularis were detected in the EEOs. Five branches were specified as having segregated from the sacral plexus: the n. cutaneus femoralis caudalis, the mutual root of n. fibularis with n. tibialis (n. ischiadicus), the rami musculares, the n. coxalis caudalis and the ramus muscularis. It was observed that the sacral plexus was linked to the lumbar plexus by the n. furcalis, to the pudendus plexus via the n. bigeminus. Consequently, the anatomic structure of the EEO's sacral plexus, the participating synsacral nerves to plexus and the innervation areas of these nerves were revealed.

Analgesic effects of intraneural injection of ethyl alcohol or formaldehyde in the palmar digital nerves of horses.

OBJECTIVE: To determine analgesic effects of intraneural injection of ethyl alcohol or formaldehyde in the palmar digital nerves of horses. ANIMALS: 6 horses. PROCEDURES: Ethyl alcohol was injected in the medial palmar digital nerve of 1 forelimb, and formaldehyde was injected in the contralateral nerve. The lateral palmar digital nerve in 1 forelimb was surgically exposed, but not injected, and the contralateral lateral palmar digital nerve was not treated. For each heel, severity of lameness in response to experimentally induced heel pain (lameness score and peak vertical force), thermal reaction time, and heel skin sensitivity scores were recorded. Heel pain was experimentally induced by advancing a threaded bolt through a custom-made horseshoe to apply pressure to the palmar horned aspect of the hoof. Horses were followed up for 112 days, when a subset of nerves was sampled for histologic analysis. RESULTS: Alcohol and formaldehyde significantly reduced all measures of heel pain, and analgesia was evident over the 112 days of the study. Pastern circumference was significantly greater for formaldehyde-treated than for alcohol-treated limbs. Histologic evaluation showed preservation of nerve fiber alignment with an intact epineurium, loss of axons, axon degeneration, fibrosis, and inflammation in alcohol-treated and formaldehyde-treated nerves. CONCLUSIONS AND CLINICAL RELEVANCE: Results suggested that intraneural injection of either ethyl alcohol or formaldehyde in the palmar digital nerves of horses resulted in substantial, but partial, heel analgesia that persisted for at least 112 days. No advantage of formaldehyde over alcohol was found, and formaldehyde resulted in greater soft tissue inflammation.

Distribution of purinergic P2X receptors in the equine digit, cervical spinal cord and dorsal root ganglia.

Purinergic pathways are considered important in pain transmission, and P2X receptors are a key part of this system which has received little attention in the horse. The aim of this study was to identify and characterise the distribution of P2X receptor subtypes in the equine digit and associated vasculature and nervous tissue, including peripheral nerves, dorsal root ganglia and cervical spinal cord, using PCR, Western blot analysis and immunohistochemistry. mRNA signal for most of the tested P2X receptor subunits (P2X1-5, 7) was detected in all sampled equine tissues, whereas P2X6 receptor subunit was predominantly expressed in the dorsal root ganglia and spinal cord. Western blot analysis validated the specificity of P2X1-3, 7 antibodies, and these were used in immunohistochemistry studies. P2X1-3, 7 receptor subunits were found in smooth muscle cells in the palmar digital artery and vein with the exception of the P2X3 subunit that was present only in the vein. However, endothelial cells in the palmar digital artery and vein were positive only for P2X2 and P2X3 receptor subunits. Neurons and nerve fibres in the peripheral and central nervous system were positive for P2X1-3 receptor subunits, whereas glial cells were positive for P2X7 and P2X1 and 2 receptor subunits. This previously unreported distribution of P2X subtypes may suggest important tissue specific roles in physiological and pathological processes.

The anatomy and physiology of the suspensory apparatus of the distal phalanx.

The equine hoof capsule protects the softer, more sensitive, structures within. Failure of the connection between hoof and bone (suspensory apparatus of the distal phalanx or SADP) results in the crippling lameness of laminitis. Active basal cell proliferation occurs principally in tubular hoof and proximal and distal lamellae. The remaining lamellae are virtually non-proliferative and the hoof wall moves past the stationary distal phalanx, by controlled activation and inhibition of constituent proteases. The lamellar corium derives most of its blood supply from the branches of the terminal arch which perforate the distal phalanx. Valveless veins within the foot can be exploited clinically for retrograde venous therapy or contrast radiography (venography). The basement membrane (BM) forms the interface between the lamellar epidermis and the adjacent dermis and the plasma membrane of each lamellar basal cell is attached to the BM by numerous electron dense adhesion plaques or hemidesmosomes the ultimate attachment unit of the SADP. Laminitis destroys and dislocates the BM and its components and without an intact, functional BM, the structure and function of the lamellar epidermis is pathologically compromised. Transcription and activation of constituent proteases occurs in normal hoof lamellae but in increased amounts during laminitis.

A pathway for predation in the brain of the barn owl (Tyto alba): projections of the gracile nucleus to the "claw area" of the rostral wulst via the dorsal thalamus.

The Wulst of birds, which is generally considered homologous with the isocortex of mammals, is an elevation on the dorsum of the telencephalon that is particularly prominent in predatory species, especially those with large, frontally placed eyes, such as owls. The Wulst, therefore, is largely visual, but a relatively small rostral portion is somatosensory in nature. In barn owls, this rostral somatosensory part of the Wulst forms a unique physical protuberance dedicated to the representation of the contralateral claw. Here we investigate whether the input to this "claw area" arises from dorsal thalamic neurons that, in turn, receive their somatosensory input from the gracile nucleus. After injections of biotinylated dextran amine into the gracile nucleus and cholera toxin B chain into the claw area, terminations from the former and retrogradely labeled neurons from the latter overlapped substantially in the thalamic nucleus dorsalis intermedius ventralis anterior. These results indicate the existence in this species of a "classical" trisynaptic somatosensory pathway from the body periphery to the telencephalic Wulst, via the dorsal thalamus, one that is likely involved in the barn owl's predatory behavior. The results are discussed in the context of somatosensory projections, primarily in this and other avian species.

[Principles of construction in the forepaw and hindpaw of the domestic cat (Felis catus). 4. Innervation of muscles and analysis of locomotion]. / Konstruktionsprinzipien an der Vorder- und Hinterpfote der Hauskatze (Felis catus). 4. Mitteilung: Muskelinnervation und Bewegungsanalysedagger.

Principles of construction in the forepaw and hindpaw of the domestic cat (Felis catus). 4. Innervation of muscles and analysis of locomotion.The innervation relations of the fore- and hindlimb of the cat were newly investigated to complete the interpretation of the muscle innervation (Figs 1-4). By means of special motion studies the contribution of paw muscles was determined during balance, locomotion as well as under specific manipulation of the prevailing single phases of the motion cycle. The functional considerations are graphically prepared in Figs 5-11.

[The most important local anesthesia in cattle: a review]. / Die wichtigsten Lokalanästhesien beim Rind: Eine Ubersicht.

Surgical interventions in cattle are frequently performed under local analgesia. Local analgesia may be carried out in the standing animal without or with slight sedation or with the animal in recumbency after deep sedation. Injection of local analgesics is less time consuming than induction and maintenance of general analgesia and is, therefore, frequently used in private veterinary practice. Precise anatomical knowledge of the nerve supply to the area to be operated is a prerequisite for the successful introduction of a local analgesia. The goal of the present review is to summarize nerve supply and indications for surgery in the area of the head, male genital tract, teat and the claws of the hind limb of cattle.

Effect of alkalinization on the local analgesic efficacy of ketamine in the abaxial sesamoid nerve block in horses.

The objective of this study was to determine the effects of the alkalinization on the local analgesic efficacy of 1% ketamine in the abaxial sesamoid nerve block in horses. Thirty-six mature healthy horses were randomly assigned to four groups for the following treatments; an abaxial sesamoid block with 5 mL of saline solution (control saline); an abaxial sesamoid block with 5 mL of a solution containing 1% ketamine (KETs 1%); an abaxial sesamoid block with 5 mL of a solution containing saline solution and 0.5 mEq of sodium bicarbonate (control bicarbonate); and an abaxial sesamoid block with 5 mL of a solution containing 1% ketamine and 0.5 mEq of sodium bicarbonate (KETb 1%). All blocks were performed in one randomly selected front leg. To determine analgesia, hoof withdrawal from thermal stimulus from radiant heat lamp was assessed. Before each block, the hoof withdrawal reflex latency (HWRL) (time between lamp illumination and withdrawal of the hoof) was determined; after the block, local analgesic effects were determined using the heat lamp at 2 and 5 min after the injection and then every 5 min for 1 h. In KETs 1% group, there were significant increases in HWRL between basal values and values from 2 to 10 min after an abaxial sesamoid block. In KETb 1% group, significant increases in HWRL was collected between the basal value and values from 2 to 25 min following an abaxial sesamoid block. In KETs 1% group, of the nine horses, four had an abaxial sesamoid block that was unsuccessful. However, in KETb 1% group, only one of the nine horses had an abaxial sesamoid nerve block that was unsuccessful. The alkalinization of a 1% ketamine solution produced a more consistent and persistent local analgesia in horses when compared with 1% ketamine solution alone.

Evaluation of the local analgesic effect of ketamine in the palmar digital nerve block at the base of the proximal sesamoid (abaxial sesamoid block) in horses.

OBJECTIVE: To evaluate the local analgesic effect of ketamine in a palmar digital nerve block at the base of the proximal sesamoid (abaxial sesamoid block) in horses. ANIMALS: 36 mature healthy Andalusian horses. PROCEDURE: Horses were randomly assigned to 4 groups of 9 horses each and received an abaxial sesamoid block in a randomly chosen forelimb with 1 of the following: saline (0.9% NaCl) solution, 1% ketamine solution, 2% ketamine solution, or 3% ketamine solution. To determine analgesia, the radiant heat lamp-hoof withdrawal model was used as a noxious thermal stimulus. Before each nerve block, baseline hoof withdrawal reflex latency (HWRL, time between lamp illumination and withdrawal of the hoof) was determined; after the nerve block, local analgesic effects were determined by measuring HWRL at 2 and 5 minutes after injection and then every 5 minutes for a total period of 1 hour. RESULTS: Significant differences in HWRL were found between baseline values and values at 2 to 15 minutes following a nerve block with ketamine. Significant differences were found between HWRL values at every time point from 2 to 10 minutes following a nerve block with saline solution, compared with 1 or 2% ketamine solution. Similarly, significant differences were found between HWRL values at every time point from 2 to 15 minutes following a nerve block with saline solution, compared with 3% ketamine solution. CONCLUSIONS AND CLINICAL RELEVANCE: Abaxial sesamoid block with ketamine ensures adequate analgesia in horses with an onset of action of 2 minutes and a maximal duration of action of 15 minutes.

Evaluation of tachykinins and their receptors to determine sensory innervation in the dorsal hoof wall and insertion of the distal sesamoidean impar ligament and deep digital flexor tendon on the distal phalanx in healthy feet of horses.

OBJECTIVE: To localize substance P (SP) and neurokinin A (NKA) and their receptors in the insertion of the distal sesamoidean impar ligament (DSIL), deep digital flexor tendon (DDFT), and dorsal hoof wall of healthy feet of horses. SAMPLE POPULATION: 18 healthy feet from horses. PROCEDURE: Samples from the dorsal hoof wall and insertion of the DSIL and DDFT of 10 feet were processed for immunocytochemical analysis, using rabbit polyclonal antisera raised against SP and NKA. Tissue sections from 8 feet were incubated with 125-labeled SP to localize tachykinin receptors and their specificity and with control solutions of radioactive SP and excess nonradioactive SP to identify areas of nonspecific binding. RESULTS: Many nerves immunoreactive for SP and NKA were localized to the region of the insertion of the DSIL and DDFT and the accompanying microvasculature and arteriovenous complexes (AVC) as well as to the microvasculature of the dorsal hoof wall. Specific neurokinin 1 receptors were localized over the microvessels and AVC of the insertion zone and small microvessels of the hoof wall. CONCLUSIONS AND CLINICAL RELEVANCE: These results document that the microvasculature of the equine foot is richly innervated and has specific receptors for tachykinins. Distributions of these tachykinin receptors on the microvasculature suggest that they form an important vasodilatory mechanism for controlling blood flow through the DSILDDFT insertion and dorsal hoof wall.

Anatomy of the distal limb.

A detailed knowledge of anatomy is essential to understand the various pathologic processes and to perform different surgical techniques involving the limbs of cattle. This article reviews macroscopic anatomy of the distal limb, muscle-tendon units, neurovascular anatomy, and external and microanatomy of the claw.

Innervation of the digit on the forepaw of the raccoon.

The innervation of the digits on the raccoon forepaw was examined by using immunochemistry for protein gene product 9.5, calcitonin-gene related peptide, substance P, neuropeptide-Y, tyrosine hydroxylase, and neurofilament protein. The larger-caliber axons in the ventral glabrous skin terminate as Pacinian corpuscles deep in the dermis, small corpuscles and Merkel endings around the base of dermal papillae, and Merkel endings on rete pegs in dermal papillae. Extensive fine-caliber innervation terminates in the epidermis and on the microvasculature. The innervation is more dense in the distal than in the proximal volar pads. Pacinian endings are also concentrated in the transverse crease separating the distal and proximal pads. In the dorsal hairy skin, hair follicles are well innervated with piloneural complexes. Merkel innervation is located under slight epidermal elevations and in some large Merkel rete pegs located at the apex of transverse skin folds just proximal to the claw. No cutaneous Ruffini corpuscles were found anywhere on the digit. The claw is affiliated with dense medial and lateral beds of Pacinian endings, bouquets of highly branched Ruffini-like endings at the transition from the distal phalanx and unmyelinated innervation in the skin around the perimeter. Encapsulated endings are located at the lateral edge of the articular surface of the distal phalanx. Extensive fine-caliber innervation is affiliated with sweat glands and with the vasculature and is especially dense at presumptive arteriovenous sphincters. Virtually all of the sweat gland and vascular innervation is peptidergic, whereas most of the unmyelinated epidermal innervation is nonpeptidergic.

The structure, innervation and location of arteriovenous anastomoses in the equine foot.

In the foot of the horse, arteriovenous anastomoses (AVAs) of epithelioid type occurred in the dermis of the coronary band, in the coronary and terminal papillae, in neurovascular bundles and at the entrance to and along the length of the dermal laminae. A particular feature of the epithelioid segment of AVAs in the horse, compared with that of other species, was the height and surface complexity of many of the endothelial cells. They extended into the lumen, forming undercut and tunnel-like areas which correlated with the characteristic surface marking of AVAs observed in vascular casts. The number of cell organelles, including the concentration of vesicles in the luminal cytoplasm, suggested cells with a high metabolic activity. The luminal surface possessed numerous microvilli and long cytoplasmic cell processes which appeared to surround material in the lumen. The innervation of AVAs was more dense than that of the arteries and consisted of adrenergic and peptidergic nerves. Noradrenaline- and neuropeptide Y-containing nerves were identified as the vasoconstrictor components of the nerve supply and occurred along arteries and formed dense plexuses around AVAs. Calcitonin gene-related peptide, substance P and vasoactive intestinal polypeptide are vasodilators and were present in single nerve fibres which accompanied arteries and AVAs along the length of the dermal laminae. In this study the distribution, density and innervation of AVAs in the equine foot are correlated with their proposed role in the development of acute laminitis. The release of vasoactive peptides from diseased organs remote from the foot may induce inappropriate prolonged dilatation of AVAs and thus contribute to the laminar ischaemia of acute laminitis.

Effects of atracurium administered by continuous intravenous infusion in halothane-anesthetized horses.

Atracurium (0.4 mg/ml in isotonic NaCl solution) was administered by IV infusion to 7 healthy adult horses for 2 hours. Over the 2-hour period, a 95 to 99% reduction of train-of-four hoof-twitch response was maintained by 0.17 +/- 0.01 mg of atracurium/kg of body weight/h, for a total of 161 +/- 6 mg of atracurium (mean +/- SEM) for horses 1 to 4, 6, and 7. Horse 5, a mare in estrus, required 0.49 mg of atracurium/kg/h to maintain comparable relaxation. Hoof-twitch recovery time from 10 to 75% of baseline strength was 19.8 +/- 2.5 minutes for all horses. The 10 to 75% recovery time for horse 5 was 18 minutes. Recovery time from discontinuation of halothane until standing was 86 +/- 14 minutes (range, 55 to 165 minutes). Horse 5 had a 165-minute recovery. Regarding recovery from anesthesia, 3 recoveries were rated as excellent, 1 recovery good, and 2 recoveries as fair. Horse 5 laid quietly until she stood with 1 strong, smooth effort.

Favored patterns in spike trains. II. Application.

In this paper we apply the two methods described in the companion paper (4) to experimentally recorded spike trains from two preparations, the crayfish claw and the cat striate cortex. Neurons in the crayfish claw control system produced favored patterns in 23 of 30 spike trains under a variety of experimental conditions. Favored patterns generally consisted of 3-7 spikes and were found to be in excess by both quantized and template methods. Spike trains from area 17 of the lightly anesthetized cat showed favored patterns in 16 of 27 cases (in quantized form). Some patterns were also found to be favored in template form; these were not as abundant in the cat data as in the crayfish data. Most firing of the cat neurons occurred at times near stimulation, and the observed patterns may represent stimulus information. Favored patterns generally contained up to 7 spikes. No obvious correlations between identified neurons or experimental conditions and the generation of favored patterns were apparent from these data in either preparation. This work adds to the existing evidence that pattern codes are available for use by the nervous system. The potential biological significance of pattern codes is discussed.

Spinal cord collaterals from axons of type II slowly adapting units in the cat.

1. The morphology of single axons, and their collaterals, of Type II slowly adapting mechanoreceptors situated at the claw bases was studied. Intra-axonal injections of horseradish peroxidase were made into the axons near their entrance to the lumbosacral spinal cord of anaesthetized cats. The morphology was revealed by subsequent histochemistry. 2. Nine Type II axons were stained. All but one bifurcated into ascending and descending branches upon entering the cord. Eighty-nine collaterals arose from the axons at a mean spacing of about 570 micrometers. 3. The collaterals formed plate-like arborizations usually about 500-600 micrometers wide in the transverse plane but only 100-300 micrometers thick in the longitudinal axis of the cord. The terminal arborizations were in laminae III-VI. 4. Synaptic boutons in laminae III and IV were more numerous than in laminae V and VI. Boutons en passant were common in laminae III and IV and arranged in series of three to six, whereas in deeper laminae only two or three boutons formed a series de passage. 5. The morphology of the slowly adapting Type II collateral is discussed. 6. Some general principles of the organization of cutaneous afferent fibres in the lumbosacral cord are presented.