Following the discovery that the proto-oncogene protein kinase B (PKB) functions as a downstream element in signalling from phosphoinositide 3′-kinase (PI3-kinase) (Burgering & Coffer, 1995), PKB has been shown to mediate a diverse array of PI3-kinase dependent cellular responses. Most recently PKB-dependent phosphorylation of 3 members of the family of Forkhead transcription factors has been demonstrated to play a role in PI3-kinase dependent effects on transcription. This review focuses on this newly discovered function of PKB in conveying the diversity of PI3-kinase dependent cellular responses.

The last decade has witnessed a significant turn in our understanding of the mechanisms responsible for the decline of cognitive functions in aged brain. As has been demonstrated by detailed morphological reassessments, the senescence-related changes in cognition cannot be attributed to a simple decrease in the number of neurons. It is becoming clearer that a major cause of age-induced deterioration of brain capability involves much subtler changes at the level of synapses. These changes are either morphological, i.e. reduction in the number of effective synapses and/or functional alterations, i.e. changes in the efficacy of remaining synapses. Important questions are now raised regarding the mechanisms which mediate these synaptic changes. Clearly, an important candidate is calcium, the cytotoxic role of which is already firmly established. The wealth of evidence collected so far regarding the changes of Ca2+ homeostasis in aged neurons shows that the overall duration of cytoplasmic Ca2+ signals becomes longer. This is the most consistent result, demonstrated on different preparations and using different techniques. What is not yet clear is the underlying mechanism, as this result could be explained either through an increased Ca2+ influx or because of a deficit in the Ca2+ buffering/clearance systems. It is conceivable that these prolonged Ca2+ signals may exert a local excitotoxic effect, removing preferentially the most active synapses. Uncovering of the role of Ca2+ in the synaptic function of the aged brain presents an exciting challenge for all those involved in the neurobiology of the senescent CNS.

Timing and pattern of expression of alkaline phosphatase was examined during early differentiation of the 1st arch skeleton in inbred C57BL/6 mice. Embryos were recovered between 10 and 18 d of gestation and staged using a detailed staging table of craniofacial development prior to histochemical examination. Expression of alkaline phosphatase is initiated at stage 20.2 in the plasma membrane of mesenchymal cells in the distal region of the first arch. Expression is strongest in osteoid (unmineralised bone matrix) and presumptive periosteum at stage 21.32. Mineralisation begins at stage E23. Expression is present in the mineralised bone matrix. Secondary cartilages form in the condylar and angular processes by stage M24. The cartilaginous cells and surrounding cells in the processes are all alkaline phosphatase-positive and surrounded by the common periosteum, suggesting that progenitor cells of the processes, dentary ramus and secondary cartilages all originate from a common pool. Nonhypertrophied chondrocytes of Meckel's cartilage express alkaline phosphatase at stage M23. Expression in these chondrocytes is preceded by the expression in their adjacent perichondrium. This is true of chondrocytes in all other cranial cartilages examined. 3-D reconstruction of expression in Meckel's cartilage also revealed that the chondrocytes of Meckel's cartilage which express alkaline phosphatase and the matrix of which undergoes mineralisation are those surrounded by the alkaline phosphatase-positive dentary ramus. By stage 25, coincident with mineralisation in the distal section of Meckel's cartilage, most chondrocytes are strongly positive. The perichondria of malleus and incus cartilages express alkaline phosphatase at stage M24. Nonhypertrophied chondrocytes along these perichondria also express alkaline phosphatase. Superficial and deep cells in the dental laminae of incisor and 1st molar teeth become alkaline phosphatase-positive at the bud stage, stages 21.16 and 21.32, respectively. Dental papillae are negative until stage M24 when alkaline phosphatase expression begins in the dental papillae and follicles of the incisor teeth and the dental follicles of the 1st molar teeth. The dental papillae of the 1st molar teeth express alkaline phosphatase at stage 25. Expression in the dental papillae and follicles appears to coincide with cellular differentiation of follicle from papilla. The presumptive squamosal, ectotympanic and gonial membrane bones, lingual oral epithelial cells connected to the dental laminae of the incisor teeth, hair follicle papillae and sheath and surrounding dermis all express alkaline phosphatase in a stage-specific manner.

Sublethal oxidative stresses increase the proportions of human fibroblasts positive for senescence associated β-galactosidase activity and accelerate the transition in the fibroblast morphotypes characterising fibroblast ageing. Stimulation of fibroblasts with TNF-α or IL-1α transiently increases the production of reactive oxygen species (ROS) in human fibroblasts. Here we propose that repeated stimulation of WI-38 fibroblasts with TNF-α or IL-1α can generate enough ROS to accelerate the transition in the fibroblast morphotypes and increase the proportion of cells positive for senescence associated β-galactosidase activity. The involvement of ROS is suggested by experiments where the stimulation of fibroblasts with TNF-α or IL-1α are performed in the presence of N-acetylcysteine which increases the intracellular antioxidant potential. It is proposed that the decrease in the proportions of morphotypes I and II, and the increase in the proportions of morphotypes III to VI observed after successive stimulation with TNF-α or IL1-α is attributed to an increased ROS production occurring during the stimulation.

The difference in the timing of the regeneration process of skeletal muscle between SJL/J and BALB/c mice was investigated using grafts of whole skeletal muscle (both autografts and allografts). Histological, autoradiographic and immunohistochemical techniques were used in the investigation. Infiltration of leucocytes into autografts, numbers of desmin-positive myogenic cells and myotube formation were all more advanced in the SJL/J compared with BALB/c mice. Furthermore, autoradiographic evidence showed that myoblasts in the SJL/J autografts were synthesising DNA 12 h earlier than myoblasts in BALB/c autografts. In allografts, where SJL/J host mice received BALB/c grafts, and vice versa, leucocyte infiltration and myotube formation occurred earlier in the BALB/c muscles grafted into SJL/J hosts, than in the reverse situation with BALB/c hosts. The results show that, at least for whole muscle grafts, it is the host environment which determines the speed and outcome of the regenerative process.

This study confirms that the median artery may persist in adult life in 2 different patterns, palmar and antebrachial, based on their vascular territory. The palmar type, which represents the embryonic pattern, is large, long and reaches the palm. The antebrachial type, which represents a partial regression of the embryonic artery is slender, short, and terminates before reaching the wrist. These 2 arterial patterns appear with a different incidence. The palmar pattern was studied in the whole sample (120 cadavers) and had an incidence of 20%, being more frequent in females than in males (1.3[ratio ]1), occurring unilaterally more often than bilaterally (4[ratio ]1) and slightly more frequently on the right than on the left (1.1[ratio ]1). The antebrachial pattern was studied in only 79 cadavers and had an incidence of 76%, being more frequent in females than in males (1.6[ratio ]1); it was commoner unilaterally than bilaterally (1.5[ratio ]1) and was again slightly more prevalent on the right than on the left (1.2[ratio ]1). The origin of the median artery was variable in both patterns. The palmar type most frequently arose from the caudal angle between the ulnar artery and its common interosseous trunk (59%). The antebrachial pattern most frequently originated from the anterior interosseous artery (55%). Other origins, for both patterns, were from the ulnar artery or from the common interosseous trunk. The median artery in the antebrachial pattern terminated in the upper third (74%) or in the distal third of the forearm (26%). However, the palmar pattern ended as the 1st, 2nd or 1st and 2nd common digital arteries (65%) or joined the superficial palmar arch (35%). The median artery passed either anterior (67%) or posterior (25%) to the anterior interosseous nerve. It pierced the median nerve in the upper third of the forearm in 41% of cases with the palmar pattern and in none of the antebrachial cases. In 1 case the artery pierced both the anterior interosseous and median nerves.

This paper presents the dynamic injury tolerances for the female humerus and forearm derived from dynamic 3-point bending tests using 22 female cadaver upper extremities. Twelve female humeri were tested at an average strain rate of 3.7±1.3%/s. The strain rates were chosen to be representative of those observed during upper extremity interaction with frontal and side airbags. The average moment to failure when mass scaled for the 5th centile female was 128±19 Nm. Using data from the in situ strain gauges during the drop tests and geometric properties obtained from pretest CT scans, an average dynamic elastic modulus for the female humerus was found to be 24.4±3.9 GPa. The injury tolerance for the forearm was determined from 10 female forearms tested at an average strain rate of 3.94±2.0%/s. Using 3 matched forearm pairs, it was determined that the forearm is 21% stronger in the supinated position (92±5 Nm) versus the pronated position (75±7 Nm). Two distinct fracture patterns were seen for the pronated and supinated groups. In the supinated position the average difference in fracture time between the radius and ulna was a negligible 0.4±0.3 ms. However, the pronated tests yielded an average difference in fracture time of 3.6±1.2 ms, with the ulna breaking before the radius in every test. This trend implies that in the pronated position, the ulna and radius are loaded independently, while in the supinated position the ulna and radius are loaded together as a combined structure. To produce a conservative injury criterion, a total of 7 female forearms were tested in the pronated position, which resulted in the forearm injury criterion of 58±12 Nm when scaled for the 5th centile female. It is anticipated that these data will provide injury reference values for the female forearm during driver air bag loading, and the female humerus during side air bag loading.

This study was undertaken to develop a functional model of otitis media with effusion (OME) in the pig (Sus scrofa), with the purpose of investigating the origin of lymphocytes populating the middle ear during the course of an inflammatory process. The relevance of the model to the human condition of OME is to a large extent dependent on the anatomical and physiological similarities between the middle ear cavity and the pharyngeal lymphoid tissue of the pig and man. Anatomical specimens were collected from 7 young Large White pigs to determine the gross anatomy of the middle ear cavity and the histological characteristics of the middle ear mucosa. It was found that the anatomy of the 3 parts of the middle ear cavity in man and in the pig is broadly similar, although some minor differences were observed. The porcine eustachian tube was seen to be cartilaginous throughout its length in contrast to the part osseous, part cartilaginous structure found in man; the porcine ossicles were slightly different in shape to those of man and the air cell system was situated inferior to the tympanic cavity in the pig as opposed to posteriorly in man. This paper describes the structure and morphology of the pig middle ear cavity and compares and contrasts it with that of man. The minor differences observed are of anatomical importance but do not diminish the usefulness of the pig middle ear cleft as a potential model for human middle ear disorders.

The lingulae of both sides of 165 dry adult human mandibles, 131 males and 34 females of Indian origin, were classified by their shape into 4 types: 1, triangular; 2, truncated; 3, nodular; and 4, assimilated. Triangular lingulae were found in 226 (68.5%) sides, truncated in 52 (15.8%), nodular in 36 (10.9%) and assimilated in 16 (4.8%) sides. Triangular lingulae were found bilaterally in 110, truncated in 23, nodular in 17 and assimilated in 7 mandibles. Of the remaining 8 mandibles with different appearances on the 2 sides, 6 had a combination of triangular and truncated and 2 had nodular and assimilated. The incidence of triangular and assimilated types in the male and female mandibles are almost equal. In the truncated type it was double in the male mandibles while the nodular type was a little less than double in the female mandibles.

The identification of myogenic precursor cells (mpc) is a key factor in determining the early events in the myogenesis and regeneration of skeletal muscle. Although satellite cells have long been established as the providers of myoblastic cells, very little is really known (apart from their anatomical location in relation to muscle fibres and their ability to migrate) about the precise role of satellite cells in myogenesis. Numerous techniques for labelling mpc have been devised, but none of these has proven to be completely reliable in firmly establishing the origin of myogenic cells. The use of tritiated thymidine to label DNA in proliferating mpc (which are not specifically distinguishable at the time) and the subsequent location of their labelled progeny in myotube nuclei has revealed a great deal of data on the timing of myogenesis, but not about the nature of mpc themselves. DNA synthesis can also be detected by antibodies to the thymidine analogue, bromodeoxyuridine, and also by antibody staining for proliferating nuclear cell antigen. Like tritiated thymidine, these other markers are not specific for muscle but are general markers for DNA synthesis. In situ hybridisation of various muscle-specific genetic markers and their products has been informative, as has immunolabelling of myogenin, MyoD1 and desmin. Desmin labelling has been particularly instructive in identifying mpc because it is one of the first muscle-specific proteins to be produced in mpc. This review covers some of the techniques mentioned above and their usefulness in determining the early events in myogenesis.

Investiture of new microvessels within an injured peripheral nerve trunk may determine the success that the local environment has in promoting axonal sprouting and regeneration. We therefore examined microvessel investment of 24 h–14 d proximal nerve stump preparations in rat sciatic nerves. The stumps, later destined to form neuromas, were created by sciatic nerve transection with resection of distal branches to prevent distal reinnervation. Microvessels were studied in the proximal stump in semithin whole mount sections of nerve and by analysis of India ink perfused microvessel profiles. Quantitative image analysis was made of the luminal profiles of vessels perfused with India ink from unfixed sections of the stumps, contralateral uninjured nerves and sham-exposed but uninjured nerves. Evidence of angiogenesis was observed in stumps 7 d after transection, indicated by a rise in the total numbers of perfused microvessels and in the numbers of 2–6 μm diameter perfused microvessels. There was a shift in the histogram of the percentage of perfused microvessels towards the 2–4 μm range and a reduction in the mean microvessel luminal area in the stumps. By 14 d, new microvessels were larger, indicated by an increase in total luminal area. New microvessels were prominent in the epineurial connective tissue or between layers of perineurial cells of former fascicles. Microvessels probably share a battery of trophic signals with other proliferating cellular elements in the milieu of the injured peripheral nerve trunk.

Several experimental models have demonstrated that the central nervous system is functionally linked to the immune system by means of the autonomic nervous system. Samples of 36 lymph nodes of patients whose ages ranged from 16 to 69 y were studied. In order to demonstrate the existence and distribution of sympathetic nerve fibres, a polyclonal antibody antityrosine hydroxylase (TH), with the streptavidin-biotin system of detection, was used. TH-positive nerve fibres appeared in all reactive patterns of the lymph nodes studied. Thin nerve fascicles ramified at the hilar region and also in the connective tissue septae. Adventitial adrenergic nerve fibres were found following afferent, and to a lesser extent, efferent blood vessels. Another source of incoming nerve fibres was found at capsular level, accompanying blood vessels. On the arterial side, the innervation ceased before reaching the follicular arterioles. Our demonstration of innervation in postcapillary venules could support a regulatory role of adrenergic neurotransmitters in lymphocyte traffic. Occasional nerve fibres were also seen in T areas among parenchymatous cells. These findings confirm the existence of sympathetic innervation in human lymph nodes, and provide indirect evidence that the psychoneuroimmune axis could also exist in humans.

The skin structure of 2 Bathyergid rodents, the naked mole-rat (Heterocephalus glaber) and the common mole-rat (Cryptomys hottentotus) is compared, to investigate whether thermoregulatory differences may be attributed to different skin features. Histological and ultrastructural studies of the dorsal skin of these closely related species show morphological and structural similarities but differences in the degree of skin folding, thickness of the integument and dermal infrastructure were evident. The skin of the common mole-rat conforms with expected morphological/histological arrangements that are commonly found in mammalian skin. Many features of the skin of the naked mole-rat, such as the lack of an insulating layer and the loosely folded morphological arrangement contribute to poikilothermic responses to changing temperatures of this mammal. Further evidence for poikilothermy in the naked mole-rat is indicated by the presence of pigment containing cells in the dermis, rather than the epidermis, as commonly occurs in homeotherms. Lack of fur is compensated by a thicker epidermal layer and a marked reduction in sweat glands. Differences in skin morphology thus contribute substantially to the different thermoregulatory abilities of the 2 Bathyergids. The skin morphology is related to the poor thermoinsulatory ability of the animals while simultaneously facilitating heat transfer from the environment to the animal by thigmothermy and/or other behavioural means.

Nuclear condensation during spermiogenesis in the ostrich follows the basic pattern established in other vertebrates. The fine granular nuclear substance of early spermatids is gradually replaced by numbers of coarse dense granules which appear to arise by aggregation of smaller dispersed elements of the chromatin. The granules increase in size and eventually coalesce to form the compact homogenous mass of chromatin typical of the mature sperm. In ostrich spermatids, however, the aggregation of the nuclear material produces large numbers of longitudinally oriented rod-shaped structures in addition to some granular material. Although fibrillar chromatin has been observed during spermiogenesis in a number of vertebrate species, the hollow nature of the rod-shaped chromatin granules in ostrich spermatids is a unique phenomenon. The spiralisation of the chromatin material observed in ostrich spermatids and in some other nonpasserine birds is possibly related to the reduction in nuclear length demonstrated during spermiogenesis in these species. In common with other nonpasserine birds, spermiogenesis in the ostrich is characterised by the appearance both of a circular and a longitudinal manchette. The circular manchette consists of a single row of microtubules reinforced by additional peripherally arranged microtubules. Links between adjacent microtubules, and between the nucleolemma and some of the microtubules, are evident. The longitudinal manchette consists of arrays of interconnected microtubules arranged in approximately 4–6 staggered, ill defined rows. This structure seems to originate as a result of the rearrangement of the microtubules of the circular manchette and is only formed once the process of chromatin condensation is well advanced. Based on the sequence of morphological events observed during spermiogenesis in the ostrich, it is concluded that the circular manchette is responsible for the initial transformation in shape of the spermatid nucleus. Thereafter, the chromatin condenses independently within the confines of the nucleolemma with the circular manchette merely acting to maintain the shape of the nucleus while this process is underway, to compress the nuclear membrane, and possibly to orientate the subunits of the condensing chromatin. The longitudinal manchette appears to assist in the translocation of material during spermatid elongation. There are indications that the developing acrosome is instrumental in effecting nuclear shaping of the apical (subacrosomal) head region of the ostrich spermatid.

The tilting angle and the shape of the glenoid cavity are considered to relate closely to shoulder stability. They are also important when planning arthroplasty and developing new designs. This study examines the glenoid cavity using 3-dimensional MRI. Forty volunteers (20 men, 20 women; average age 21.4; range 18–35 y) were enrolled in the study. The tilting angles of the glenoid bone were measured in 5 consecutive axial planes perpendicular to the glenoidal long axis. Cross sections were divided into 3 types (concave, flat, convex) according to the shape on each plane.

The average tilting angles for the 5 planes from the bottom to the top were 3.3±4.1, 1.4±3.8, −0.6±1.9, −1.4±3.3, and −6.2±3.3 degrees anteriorly, indicating that the 3-dimensional bony structure of the glenoid was twisted anteriorly to posteriorly. Images on the bottom plane consisted of 82.5% concave type, 15% flat type and 2.5% convex type, while only 3 cases (7.5%) showed concave at the top plane. The shape of the glenoid cavity is thought to be conducive to glenohumeral motion and stability.

The mouse thymus changes dramatically during pregnancy. It shrinks in size, and the cortex is extensively reduced from midpregnancy onwards. Despite this, there is surprisingly little evidence for any increase in apoptosis, and considerable evidence that mitosis of thymocytes continues throughout pregnancy. In spite of overall involution the thymic medulla actually expands in midpregnancy due to a combination of mitosis of epithelial cells and an accumulation of lymphocytes. The extent and nature of these changes are examined in this study at the ultrastructural level. The epithelial cells of the subcapsular cortex (type 1 cells) become wrinkled and exhibit powers of phagocytosis, whilst the other cortical epithelial cells are relatively unchanged, although the formation of epithelial/thymocyte rosettes and thymic nurse cells is more clearly seen in midpregnancy than usual. Other changes associated with pregnancy involve the medullary epithelial cells that undergo an increased level of mitosis. Their greater numbers surround accumulations of lymphocytes to form the characteristic medullary epithelial rings. Cell movement through blood vessel walls was clearly observed in midpregnancy, but not at other times. Interdigitating cells in the medulla become more conspicuous as pregnancy proceeds and the cells become phagocytic. The endoplasmic reticulum in plasma cells becomes expanded, indicating increased secretory activity. These results highlight the active nature of the thymus in pregnancy in spite of its involution. This picture contradicts the conventional notion that an involuted thymus is inactive.

The accessibility of the primary sensory neurons of the trigeminal system at stages throughout their development in avian and mammalian embryos and the ease with which these neurons can be studied in vivo has facilitated investigation of several fundamental aspects of neurotrophin biology. Studies of the timing and sequence of action of neurotrophins and the expression of neurotrophins and their receptors in this well characterised neuronal system have led to a detailed understanding of the functions of neurotrophins in neuronal development. The concepts of neurotrophin independent survival, neurotrophin switching and neurotrophin cooperativity have largely arisen from work on the trigeminal system. Moreover, in vitro studies of trigeminal neurons provided some of the first evidence that the neurotrophin requirements of sensory neurons are related to sensory modality. The developing trigeminal system has been studied most extensively in mice and chickens, each of which has particular advantages for understanding different aspects of neurotrophin biology. In this review, I will outline these advantages and describe some of the main findings that have arisen from this work.

The lacrimal sac and nasolacrimal duct are surrounded by a wide cavernous system of veins and arteries comparable to a cavernous body. The present study aimed to demonstrate the ultrastructure of the nervous tissue and the localisation of neuropeptides involved in the innervation of the cavernous body, a topic not previously investigated. Different S-100 protein antisera, neuronal markers (neuron-specific enolase, anti- 200 kDa neurofilament), neuropeptides (substance P, neuropeptide Y, calcitonin gene-related peptide, vasoactive intestinal polypeptide) and the neuronal enzyme tyrosine hydroxylase were used to demonstrate the distribution pattern of the nervous tissue. The ultrastructure of the innervating nerve fibres was also examined by means of standard transmission electron microscopy.

The cavernous body contained specialised arteries and veins known as barrier arteries, capacitance veins, and throttle veins. Perivascularly, the tissue was rich in myelinated and unmyelinated nerve fibres in a plexus-like network. Small seromucous glands found in the region of the fundus of the lacrimal sac were contacted by nerve fibres forming a plexus around their alveoli. Many nerve fibres were positive for S-100 protein (S 100), neuron-specific enolase (NSE), anti-200 kDa neurofilament (RT 97), calcitonin gene-related peptide (CGRP), substance P (SP), tyrosine hydroxylase (TH), and neuropeptide Y (NPY). Vasoactive intestinal polypeptide (VIP) immunoreactivity was only demonstrated adjacent to the seromucous glands.

Both the density of nerve fibres as well as the presence of various neuropeptides emphasises the neural control of the cavernous body of the human efferent tear ducts. By means of this innervation, the specialised blood vessels permit regulation of blood flow by opening and closing the lumen of the lacrimal passage as effected by the engorgement and subsidence of the cavernous body, at the same time regulating tear outflow. Related functions such as a role in the occurrence of epiphora related to emotional responses are relevant. Moreover, malfunction in the innervation of the cavernous body may lead to disturbances in the tear outflow cycle, ocular congestion or total occlusion of the lacrimal passages.

It is currently thought that New World monkeys, prosimians, and humans are the only primates to possess vomeronasal organs (VNOs) as adults. Recent studies of the human VNO suggest that previous investigations on Old World primates may have missed the VNO. We examined nasal septa from the chimpanzee (Pan troglodytes) grossly and histologically for comparison with nasal septa from humans, Old World monkeys (Macaca fascicularis, M. nemistrina) and prosimian primates (Microcebus murinus, Otolemur garnettii). Grossly, chimpanzees had depressions on the nasal septum similar to fossae reported anterior to the VNO openings in humans. Histologically, chimpanzees and humans had bilateral epithelial tubes which were above the superior margin of the paraseptal cartilages (vomeronasal cartilage homologue). The epithelial tubes had a homogeneous ciliated epithelium. These structures were thus positionally and structurally identical to the human VNO and unlike the well-developed prosimian VNOs which were surrounded by vomeronasal cartilage. Macaques had no structures which resembled the VNO of either the prosimians or humans. The results demonstrate that the VNO is present postnatally in the chimpanzee and is almost identical to the human VNO in its anatomical position and histological structure. This in turn suggests that the reported absence of the VNO in at least some adult Old World primates is artifactual, and that further study may provide evidence for its existence in other species.

The arrangement of the enteric ganglia and nerve fibre plexuses was examined in the submucous and mucous layers and around Peyer's patches of the porcine small intestine to clarify their organisation. Immunohistochemistry of vasoactive intestinal peptide (VIP) and neurofilament proteins in wholemounts, chopped or paraffin sections was used to locate the neural elements. The ganglia of the internal and external submucous plexuses were situated at 2 different topographic locations, being clearly demarcated by the submucosal vascular arcades and differing in neuronal composition. The internal submucous plexus was the only contributor to the plexus surrounding the follicles of Peyer's patches as a continuous mesh of 3 ganglionated nerve subplexuses. VIP-immunoreactive fibres from this mesh innervated the dome. The mucosal plexus, which was subdivided into 4 subunits — the outer proprial, inner proprial, pericryptal and villous plexuses — contained a few solitary neuronal perikarya. Labelling for neurofilament proteins revealed Dogiel types II, IV and VI neurons. The observations reveal several new features in the enteric nervous system of the pig and clarify its nomenclature.