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A new morphological type of somatosensory organ is described. It is found in the glabrous skin of the dog nose (rhinarium or planum nasale) and situated in dermis papillae. The otherwise thick epidermis forms a thin window above the organ. There are only a few layers of keratinocytes in the window and the corneocytes are much thinner than elsewhere. The organ consists of highly branching cells that wrap naked nerve endings emanating from myelinated nerve fibers originating in the outer dermal nerve plexus. The structure entirely fills the top of the dermal papilla. The intercellular spaces of the organ and its surroundings are occupied by an extended areolar basal lamina., (© 2020 The Authors. Journal of Morphology published by Wiley Periodicals, Inc.)

The usual pigmentation pattern in mammalian skin consists of fixed melanocytes in the basal layer of the epidermis, supplying keratinocytes with melanosomes. We observed that the glabrous skin (rhinaria and footpads) of dogs deviates from this pattern. In dogs, melanocytes are found in both the dermis and epidermis. The epidermal melanocytes are situated in the intercellular spaces of the basal and spinous layers. They are characterized by a quantity of cytoplasm containing a centriole, also developing melanosomes, and in some cases annulate lamellae. There is a high frequency of closely apposed melanocytes in the epidermis. Melanosomes in different stages of formation are also abundant. The morphology of the glabrous skin of dogs suggests transport of melanocytes from the dermis into the epidermis and formation of melanosomes in the epidermis. A distributed and intense pigment formation may be necessary to achieve the black noses of many dog breeds and wild canids, as well as dark footpads despite heavy abrasion and rapid skin renewal., (© 2018 Wiley Periodicals, Inc.)

The skin of the lemur nose tip (rhinarium) has arterioles in the outer vascular plexus that are endowed with an unusual coat of smooth muscle cells. Comparison with the arterioles of the same area in a number of unrelated mammalians shows that the lemur pattern is unique. The vascular smooth muscle cells belong to the synthetic type. The function of synthetic smooth muscles around the terminal vessels in the lemur rhinarium is unclear but may have additional functions beyond regulation of vessel diameter.

Most mammals have nose tips covered by glabrous skin, a labronasal area, or rhinarium. The surface of the rhinarium of Lemur catta has a dermatoglyphic pattern consisting of epidermal domes. Below the domes, epidermal pegs dip down into the dermis. In and below the tip of the epidermal peg, a complex sensory organ is found. It consists of an association of innervated Merkel cells, lamellate (Pacini-like) bodies with a central nerve, and a ring of unmyelinated nerve endings in the epidermis. The Merkel cells are situated basally in the epidermis and the lamellated bodies just below the epidermis. The unmyelinated nerve endings related to the organ ascend in a circle straight through the epidermis ending below the corneal layer. From these nerve terminals, horizontal spikes enter the keratinocytes. The three components occur together forming an organ and are innervated from a common nerve plexus. The morphology of the complex sensory organ of the lemur shares most crucial components with Eimer's organs in moles, echidna, and platypus, while some structures are lacking, for example, the specific central pillar of keratinocytes, the cuticular cap, and a central unmyelinated fiber. The presence of the essentials of an Eimer's organ in many mammals suggests that a wider definition is motivated., (© 2015 Wiley Periodicals, Inc.)

The striated muscles of Derocheilocaris typica consist of mononucleated cells, each containing one filament bundle. Large muscles consist of two or more cells adjacent to each other. The mitochondria line up along the filament bundle on one side. The nucleus is situated in the mitochondrial row and has a small cytoplasmic area around it filled with glycogen. The sarcomeres are between 3 and 6 μm long. The Z-line and H band are present. Six thin filaments surround one thick filament. All muscles belong to the phasic type. The tubular system emanates from the ends of the muscle cell and penetrates the whole cell. The tubules are formed as cisterns, which also open at the cell membrane at the level of the I bands. They have sarcoplasmic cisterns on both sides forming a continuous triad system. Partially transformed epidermal cells mediate muscle insertions on the cuticle. Tendons are formed with the transformed epidermal cells being supplemented by fibroblasts forming collagen fibers. Dorsal and ventral abdominal muscles are innervated from the dorso-lateral nerve arising from the nerve chain. Each muscle cell receives one axon, which forms one synapse on the mitochondrial-free side of the muscles. Axons form terminal spines, which make axo-axonal synapses., (Copyright © 2013 Elsevier Ltd. All rights reserved.)

Mystacocarids are dioecious. Their gonopores are on the medial side of the third thoracic limb. The male's paired testes lie in the thorax and abdomen. They develop from paired rows of six small follicles dorsally. In the mature animal they fill most of the abdomen. The spermatophores develop within the follicles from spermatogonia mixed with follicle cells, which support and nourish the spermatocytes and produce the seminal fluid. The short vas deferens runs along the bottom of the testes and then continues forward to the gonopore. The vas deferens has a small group of cells near the gonopore that becomes a closure mechanism. The female has reproductive cells and also support cells that provide nutrition and form the wall of the ovary and oviduct. The unpaired female ovary begins in the third thoracic segment. During maturation, the oocytes are pushed posteriorly. The enormous mature ovum extends into a caudal pocket of the ovary. Starting with its anterior end, this ovum is extruded into the short oviduct, which extends laterally and ventrally to the gonopore. During extrusion, the pocket is reabsorbed from behind. There are no accessory structures connected to the reproductive system, nor any external specializations on the third limb., (Copyright © 2012 Elsevier Ltd. All rights reserved.)

The world-famous 'Orsten' Konservat-Lagerstätte has yielded detailed information about Cambrian arthropods and their morphology. Internal organs or soft tissues have, however, rarely been reported, an obvious palaeobiological drawback. In this study, we employed synchrotron radiation X-ray tomographic microscopy (SRXTM) to study microscopic 'Orsten' arthropods from the Cambrian of Sweden: Skara minuta and two phosphatocopine species, Hesslandona sp. and Hesslandona trituberculata. This exceptionally high-resolution technique reveals internal organs or soft tissues that allow detailed comparison with equivalent structures in extant crustaceans and functional inferences to be made. The S. minuta specimen shows the digestive system and muscles that extend to the extremities. The slanting anterior portion of the head and anterior position of the mouth with a straight oesophagus suggest a primarily brushing and scraping way of feeding. The prominent head appendage muscles indicate muscle strength and good capacity for food manipulation. In the phosphatocopines the bulbous labrum is one of the most prominent morphological structures of the body. All specimens analysed reveal pairs of muscle bundles within the labrum. Based on comparisons with extant crustacean relatives, these muscles would fulfil the function of moving the labrum up and down in order to open the buccal cavity. The results of this pilot study demonstrate that there is still much to be learned about the 'Orsten' taxa.

An ultrastructural study of the intestine of Derocheilocaris typica revealed an organization of the midgut musculature, which is unique in the Crustacea. This species unusual anal skeletomusculature has also not been seen before. The intestinal musculature of D. typica displays different patterns in the fore-, mid-, and hindgut. Around the foregut, eight pairs of dilator muscles complement a contiguous carpet of circular muscles around the foregut. Their coordinated action serves to suck in food and pass it to the midgut. A pair of large glands, each consisting of three cells, opens into the foregut above the mouth. The midgut musculature differs from any previously described. Circular muscles give rise to thin, longitudinal protrusions and short longitudinal muscles. The distribution of all of them is irregular. Thus the short longitudinal muscles, which have a length of approximately one segment, vary from none to five within a segment. The last abdominal segment is exceptional, by having 15-20 short longitudinal muscles. The hindgut has three longitudinal muscle groups each consisting of three muscles, one dorsally and one on each side. The posterior end of the midgut and the hindgut suggests that they act together to achieve defecation. The importance of the peri-intestinal cells as part of the nutritional process is emphasized., (Copyright 2010 Elsevier Ltd. All rights reserved.)

The mystacocarid crustacean Derocheilocaris typica has two microvillar organs, one new, the other previously unappreciated in crustacean literature. The first is situated on the head-shield and consists of three pairs of cells: one with microvilli and a ballooned nucleus; one smaller and without special features; the third large and investing the other two and extending down to the foregut. We call this new organ the "cephalic microvillar organ" and discuss the value of the concept "dorsal organ", to which it might have been included. The second organ consists of about 21 cells that cover the proximal part of the dorsal surface of the labrum. The cells are alike, being characterized by an apical field of microvilli and a large residual body. This organ is here called the "labral microvillar organ". Both organs are neither sensory nor secretory and do not qualify for membership in any of the other recognized organ systems. We are unable to deduce their Dero-cheilocaris functions.

An excretory antennal gland, composed of only eight cells, is found entirely in the limb in the mystacocarid Derocheilocaris typica. The end sac is composed of podocytes, valve cells and cap cells. The podocytes contain enormous residual vesicles. There are few pedicel complexes, and they arise directly from the cell surface without intermediate foot processes. The excretory duct is entirely lined with microvilli, which are separated from the lumen by a modified layer of thin cuticle.

Frontal eyes of crustaceans (previously called nauplius eye and frontal organs) are usually simple eyes that send their axons to a medial brain centre in the anterior margin of the protocerebrum. Investigations of a large number of recent species within all major groups of the Crustacea have disclosed four kinds of frontal eyes correlated with taxonomic groups and named after them as the malacostracan, ostracod-maxillopodan, anostracan, and phyllopodan frontal eyes. The different kinds of eyes have been established using the homology concept coined by Owen [Owen, R., 1843. Lectures on the comparative anatomy and physiology of the invertebrate animals. Longman, Brown, Green, Longmans, London] and the criteria for homology recommended by Remane [Remane, A., 1956. Die Grundlagen des natürlichen Systems, der vergleichenden Anatomie und der Phylogenetik. 2nd ed. Akademische Verlagsgesellschaft, Geest und Portig, Leipzig]. Common descent is not used as a homology criterion. Frontal eyes bear no resemblance to compound eyes and in the absence of compound eyes, as in the ostracod-maxillopodan group, frontal eyes develop into complicated mirror, lens-mirror, and scanning eyes. Developmental studies demonstrate widely different ways to produce frontal eyes in phyllopods and malacostracans. As a result of the studies of recent frontal eyes in crustaceans, it is concluded by extrapolation that in crustacean ancestors four non-homologous frontal eye types evolved that have remained functional in spite of concurrent compound eyes.

Immunocytochemical techniques for double staining were employed to investigate the morphological basis for interactions between enkephalins and other neuroactive compounds in the behavior of the gastropod mollusc Cepaea nemoralis. Coexistence of each of the two enkephalins with FMRFamide, serotonin or GABA-like immunoreactivity was found in certain neurons in cerebral, parietal, and pedal ganglia. Tyrosine hydroxylase-immunoreactive neurons were occasionally seen in close apposition to, but never colocalized with, the enkephalins. A comparison between these anatomical observations and previous behavioral studies suggests that in gastropod molluscs cotransmission of enkephalins with classical transmitters may, at least partly, reflect synergism of these substances in the control of definite behavioral programs.

NADPH-diaphorase (NADPHd) is known to be identical to nitric oxide (NO) synthase in the mammalian nervous system, and is therefore used as a marker of NO-producing neurones. Using the histochemical reaction for NADPHd, we searched for such neurones in a selection of invertebrates. Special emphasis was given to molluscs. No selective neuronal staining was found in representatives of coelenterates, turbellarians, nematodes and urochordates. In all annelids, arthropods and molluscs examined, with the exception of a chiton, specific neurones were selectively stained. The reaction was particularly strong in pulmonate molluscs where scattered positive neurones were found in various ganglia and clustered symmetrically in the paired buccal ganglia. Biochemical assay of NO synthase in osphradia of the gastropod mollusc Lymnaea stagnalis revealed a formation of citrullin that was inhibited by the specific NO synthase N omega-nitro-L-arginine (NO2Arg). Both histochemical and biochemical methods indicate that NO can be used as a signal molecule by specific neurones in advanced invertebrates.

The structure of the aesthetascs has been investigated in the prawn Macrobrachium rosenbergii (larvae and juveniles), the opossum shrimp Neomysis integer, the euphausid Meganyctiphanes, and in the water-fleas Daphnia magna and D. longispina. The aesthetascs, that are thought to represent olfactory receptors, exhibit a considerable structural variation, ranging from the well known aesthetascs of higher crustaceans (lobster, crab, crayfish) to the corresponding sensilla found in the water-fleas and the males of opossum shrimps. The two following morphological characteristics of the aesthetascs are thought to indicate an olfactory function: the shape of the cuticular hair that is long and essentially hose-shaped, and the thin, loosely arranged cuticle of at least the outer part of the cuticular hair. The presence of other structural elements such as sensory cells, cilia, and enveloping cells are vital for the olfactory function, but the development is variable, which makes their use in the morphological definition of aesthetascs problematic.

Each of the approximately 500 ommatidia in the compound eye of the cladoceran crustacean Leptodora kindtii has a crystalline cone consisting of five cells. Five retinula cells are also present, one of which contributes to the distal 1-2 micrometer of the rhabdom only; the other four retinula cells from a continuous rhabdom. Throughout the rhabdom its cross section displays two separate halves with the axis of the microvilli in one half perpendicular to that in the other (orthogonal pattern). Interferometric analysis of the refractive index of the crystalline cone revealed an inhomogeneous system with one distal and one proximal gradient. The gradient system was found to exclude rays entering from adjacent facets, thus maintaining the optical isolation. Consequently, these optics replace distal screening pigment, which is absent in the eye. The long and unscreened crystalline cones give rise to an almost transparent eye in conformity with the overall transparency of this plank-tonic animal. The morphological characteristics of the eye of this species deviate from other cladoceran eyes, but the optical design closely resembles that of some pelagic marine amphipod crustaceans.

Serial sections of human vaginal and keratinized oral-gingival epithelia were investigated for ciliary structures. Most melanocytes of the gingival epithelium lacked cilia, whereas almost all basal keratinocytes of the deeper portion of the epithelial ridges possessed one cilium each. In the suprabasal layers of the ridges only a few keratinocytes exhibited a single cilium. In the basal layer, at the top of the connective tissue papillae, approximately every second keratinocyte displayed a single cilium. In the suprabasal layers above the ridges no ciliated keratinocytes were observed. The basal cells of the vaginal epithelium were endowed with cilia, while cilia were absent from the suprabasal cells. In the human forearm epidermis most melanocytes and keratinocytes are supplied with a single cilium; it has been suggested that they may play a role in light reception. However, the widespread occurrence of 9 + 0 cilia in epithelial cells of internal epithelia and their coincidence with the sites of renewal of keratinocytes suggests that a relationship may exist between solitary cilia and mitotic activity.

Human keratinocytes were investigated for the presence of single cilia. Almost all basal keratinocytes were found to carry a single cilium in normal, occluded, and psoriatic skin. The ciliary structure was progressively reduced in keratinocytes approaching the surface. No remnants of the ciliary apparatus were found in the granular layer. In one case of nickel-allergic dermatitis (patch test), the keratinocytes had lost their cilia; the significance of this surprising finding remains to be elucidated.

3,4-dihydroxyphenylalanine (DOPA), 3,4,5-trihydroxyphenylalanine (5-OH-DOPA), 5-S-cysteinylDOPA (5-SC.D) and 2-S-cysteinylDOPA (2-SC.D) in the tentacles of the sea anemone, Metridium senile, were studied by the combined use of differential centrifugation of tissue homogenates, ultracentrifugation on sucrose density gradients, HPLC and electron microscopy. DOPA, 5-OH-DOPA and o-diphenol:O(2) oxidoreductase (Tyrosinase) were enriched in fractions containing membranes and subcellular particles, and the cysteinylDOPAs in the cytosol fractions. Ultrastructurally studied fractions rich in DOPA and 5-OH-DOPA contained large numbers of highly osmium-reducing vesicles. Identical structures were localized in ectodermal nerves and epidermal sensory cells. The results suggest that previous findings of catecholderivatives in the tentacles of Metridium and other sea anemone species by histochemical methods, are explained by a tyrosinase-based accumulation of DOPA and 5-OH-DOPA in the ectodermal nerve net. These substances are confined in specific compartments (vesicles) in the neurons and sensory cells.

The crustacean optic neuropiles, the lamina ganglionaris and especially the medulla externa, show a specific pattern of green fluorescence with the fluorescence histochemical method of Falck-Hillarp. Normally, only the terminals and the cell bodies fluoresce, but in reserpine-treated animals exogenous catecholamines are taken up by the whole adrenergic neuron and are thus visualized as a whole. Incubating crayfish optic neuropiles in dopamine or alpha-methylnoradrenaline after reserpine treatment demonstrated a tangential neuron connecting the lamina and the medulla externa. The morphology of this tangential neuron differs from the two types of tangential neurons, Tan1 and Tan2, previously characterized with Golgi techniques. The catecholaminergic neuron thus constitutes, a third tangential neuron type.

Certain neurons in the nervous system of the malacostracan crustaceans give rise to a predominantly green and a sparse yellow fluorophore in the histochemical fluorescence method of Falck-Hillarp. The same applies to the whole of Crustacea. The green fluorophore is probably a catecholamine; the yellow to brown-yellow has not yet been identified. The biogenic amine responsible for the green fluorescence, besides being found in diffusely distributed fibres, also appears in distinct areas of fibre concentrations in the central nervous system. The protocerebrum of the malacostracans contains three areas: the central body and two areas in the top of the brain, one anterior and one posterior. The latter two are not recognized as separate areas in ordinary histological preparations. In addition, the optic neuropiles are fluorescent, some with a distinct stratification of the fluorophore. The deuto- and tritocerebrum and the ventral nerve cord also contain monoaminergic neurons. Of the brightly fluorescent areas in the whole of Crustacea, only the central body consistently exists in all species. The other areas of concentrated fluorescent neuropile are restricted to smaller taxonomic units and differ from each other.

The isopod species Astacilla longicornis (Sowerby) has a layered rhabdom, which is the first case reported in isopod crustaceans. The rhabdom comprises the rhabdomeres of six retinular cells. It is surrounded by an extracellular palisade. Usually, palisades are intracellular in arthropods. A layered rhabdom is found in taxonomically widely separated groups within the Arthropoda and no intermediate forms are found between the different, highly specialized rhabdom types present. It is concluded that the layered rhabdom represents an example of convergent development.

Antiserum against bovine retinal S-antigen, a soluble protein (MW = 50 kDa) thought to be involved in phototransduction, was used in an immunohistochemical and immunochemical study of vertebrate eyes and pineal systems and invertebrate photoreceptor organs. Positive reactions, not seen with antiserum preabsorbed with highly purified S-antigen, were observed in planarian and starfish ocelli; scallop eyes; polychaete eye; crayfish compound eye; lamprey, salmon, frog, turtle, quail and hamster eyes. A specific reaction was also seen in the pineal organ of all the vertebrates examined, albeit weak in turtle and quail. In addition, several structures associated with photoreceptor organs, including the reduced frontal eyes of crayfish, the organ of Bellonci in crayfish eyestalk, and bipolar cells resembling those giving rise to Landolt's clubs in quail and golden hamster retinae, were immunopositive. Immunochemical studies revealed the presence of a single immunopositive band of protein which was similar but not identical in size in all vertebrate eyes and pineal organs (except that of chicken pineal) and invertebrate tissue examined. The wide distribution of positive reaction in photoreceptive tissue indicates that the retinal S-antigen determinant has been highly conserved during evolution.

The Langerhans cells are capable of taking up L-dopa and the catecholamines dopamine and noradrenaline when exposed to these substances in vitro. Within the cell L-dopa is found in the cytoplasm as well as in the nucleus, whereas the catecholamines are confined to cytoplasmic granules. The L-dopa uptake is most probably carrier-mediated and the hypothesis is brought forward that L-dopa enters the cell by exchange diffusion. At present little is known about the nature of the amine uptake mechanism.

A comparative investigation of the distribution of monoaminergic neurons in non-malacostracan crustaceans was performed with the histochemical fluorescence method of Falck-Hillarp. Two fluorophores were found: the more widespread of the two emits a green fluorescence; and the more sparsely distributed emits a yellow to brown-yellow fluorescence. Specific green fluorescent areas were shown to exist in the protocerebrum. The central body and the optic ganglia of the compound eye (where present) are always fluorescent. Moreover, the centre of the nauplius eye may have a green fluorophore, as in ostracods, and a neuropile area, here called the frontal area. These neuropile centres are known from ordinary histological studies of the nervous system. In addition, there are specific monoaminergic centres, such as the so-called dorsal area of phyllopods and anostracans as well as the copepod specific areas. Specific monoaminergic areas appear in the deutocerebrum and the suboesophageal ganglion where they are particularly well developed. Presumed sensory neurons in the cavity receptor organ of Artemia saliva are shown to be monoaminergic. Monoaminergic sensory neurons have not been described previously in Arthropods. Presumed motor innervation of hind-gut and trunk muscles is also found, and it is concluded that in crustaceans neurons of every type (sensory, internuncial, motor) may be monoaminergic.

The neuronal types and patterns in the visual system of the species Artemia salina and Daphina magna have been studied with the Golgi method and electron microscopy. The lamina contains five classes of neurons: photoreceptor axons, monopolar, centrifugal, tangential and amacrine neurons. The terminals of the receptor axons are distributed in two (A. salina) or three (D. magna) layers. The dilated terminals have an extensive and wide array of fine branches. One axon from each ommatidium bypasses the lamina and terminates in the medula in A. salina. A. salina has four types of monopolar neurons, two of which are stratified, whereas in D. magna only two types are found, one of which is bistratified. Tangential T-neurons connect the lamina with the protocerebrum. D. magna has in addition one tangential T-neuron connecting both the lamina and the medulla with the protocerebrum. In both species monopolar-type centrifugal neurons connect the medulla and the lamina, whereas that of A. salina has a wide laminar distribution. Both species also have amacrine cells in the lamina. The medulla contains, besides those shared with the lamina, transmedullary neurons (two types in A. salina), amacrine cells and neurons originating in the protocerebrum. "Cartridge"-type synaptic compartments are lacking in the investigated species, although a periodic arrangement is discernible in the distal portion of the lamina of A. salina. The receptors from three types of specialized contacts in Artemia, one of which involves a dyad. D. magna has only one-to-one synapses. Neurosecretory fibres are absent in A. salina.

The crustacean species Pacifastacus leniusculus and Gammarus pulex were investigated by electron microscopy in a search for possible neuromuscular junctions in the hindgut, which has a rich supply of catecholaminergic fibres. True neuromuscular synapses were found in both species between nerve terminals containing dense-core vesicles (80-110 nm in diam.) and muscle fibres. We suggest that the dense-core vesicle terminals contain a catecholamine, and this is supported by ultrahistochemical tests for monoamines. Two types of junctions are found: one in which the nerve terminal is embedded in the muscle cell (both species) and one in which protrusions from the muscle cell meet nerve terminals (Pacifastacus). Gammarus pulex, which has only circular muscles in the hindgut, has only catecholaminergic innervation, whereas Pacifastacus leniusculus has circular and longitudinal muscles both with at least two types of innervation.

We have investigated, comparatively, the ontogenetic development of the compound eye in larvae of a mysid (Neomysis) and a euphausiid (Thysanoessa) species and found it to be close to identical in the two species. The larval eye is of apposition type with special adaptations for planktonic life. The elongated dioptric apparatus is devoid of screening pigment and instead has a proximal lens optically isolating the ommatidium. The pigmented retina is extremely compressed making the eye largely transparent and presumably suitable for a planktonic life. The presence of this specialized type of eye in the planktonic larvae of euphausiids was known before but it is intriguing to find exactly the same type in mysids, spending their entire larval life as embryos in the female marsupium. A possible explanation is offered if mysids earlier in evolution had planktonic larvae. Upon reduction of free-living larvae, the transparent type of eye may have been preserved because there is no selection pressure on the larva to change it. In late larval life, both species transform their eyes to a refracting superposition type typical for adult mysids and euphausiids. The process of transformation and the functional connection between transparent apposition and superposition is described.