The connective tissue skeleton in the mammalian kidney and its innervation.

A special reticular basket-like system of connective tissue strips (called connective tissue skeleton - CTS) was found between the cortex and medulla in the kidney of various mammals. It enlarges the wall of the renal calyx (or pelvis) into the parenchyma. The main component of this system is collagen. A small amount of smooth muscle cells was found in one part of CTS srips situated around the papilla (the levator fornicis muscle). A dense monoaminergic and scanty cholinergic innervation was found in the whole system of the CTS. The functional importance of this system is discussed: (i) a tightly linked connection between the urine-discharging system and the kidney, (ii) 'milking' and similar effects in the papilla as well as perception of intrapelvic and intrarenal pressure, (iii) penetration of infection into the kidney and (iv) liberation of the monoaminergic transmitter.     

Kidney of the great Indian rhino Rhinoceros unicornis, Linnaeus

The kidney of R. unicornis has almost 80 closely apposed lobes, all appearing peripherally. Every lobe, almost enclosed by a collagenous septum, resembles a deformed truncated cone. The pelvis proper is a small pouch which divides into a cephalic and a caudal urothelial-lined fibromuscular conduit. The terminal collecting ducts of every lobe open into a tubus maximus. This is lined by cuboidal cells and otherwise has no wall. There is no papilla. All lobes finally empty through the 18 primary infundibular orifices at the pelvic conduits. A primary fibromuscular infundibulum typically yields a secondary one supplying an adjacent lobe. Two or three lobes can use a common tubus maximus by "convergence" of their medullae. Tubus maximus, terminal collecting ducts and deep outer medulla are embraced by a fibromuscular calyx which is the peripheral extension of an infundibulum and is fused to the outer medulla. There is thus no vault between medulla and calyx. Large intralobar veins are fused to the outer wall of the calyx. The possible significance of this is discussed. The cortex is the only part of a lobe which has contact with infundibulum, pelvic conduits, or pelvis proper. The kidney has about 16 million glomeruli which form 5.8% of the adult's cortical mass. Many adult mammals, from mouse to rhinoceros, fit into the log10-log10 slope relating number of glomeruli per kidney to body-mass. Neonatal rhinos at term have mature glomeruli throughout the cortex. The small size of the glomeruli and the large number per field allow 16 million in an 118-gm kidney.     

Comparative study of the topographic localization of NADPH diaphorase positive cells in the rabbit and pheasant thymuses

Nicotinamide adenine dinucleotide hydrogen phosphatediaphorase (NADPHd) histochemistry was used as a marker for nitric oxide synthase (NOS). In the rabbit thymus, NADPHd staining was observed between the capsule and corticomedullary junction in radially oriented blood vessels in the cortex. The outer surface of the thymic lobule and interlobular septa showed adipocytes clumped together. There was a high density of NADPHd positive cells in the medulla, without a sharp boundary in corticomedullary space. In addition to radially oriented blood vessels in the cortex, they were also found as solitary profiles with well stained walls in the medulla. Neuronal plexuses were localized in perivascular topography. In the pheasant thymus, NADPHd positive cells were present as clusters which were distributed in the medulla and the corticomedullar area. NADPHd positive nerve fibres were localized in perivascular topography.     

An experimental model of ascending pyelonephritis due toCandida albicans in rats

We developed a new experimental model of ascendingCandida pyelonephritis in female rats with leukopenia and vesicoureteral reflux. Rats were treated transperitoneally with cyclophosphamide (200 mg/kg) to induce leukopenia 3 days before and transurethrally with diluted acetic acid solution to induce vesicoureteral reflux 1 day before inoculation ofCandida albicans strain, ATCC 10259 (containing 10⁷ cells). Microscopy revealed acute pyelonephritis in whichCandida cells invaded from the fornix and/or papilla into the medulla within 3 days after inoculation. Between 7 and 28 days after inoculation, chronic pyelonephritis reached the cortex. The incidence of pyelonephritis increased gradually and was approximately 80% after 7 days.Candida colony counts of bladder urine specimens obtained by direct puncture were significantly greater in rats with pyelonephritis extending into the parenchyma than in those with pyelonephritis located along the pelvis (p<0.01). These results suggest that this rat model shows the characteristic feature of ascending pyelonephritis due toC. albicans and that the severity ofCandida pyelonephritis can be estimated fromCandida counts of bladder urine.     

A scanning electron-microscopic study of the rat thymus with special reference to cell types and migration of lymphocytes into the general circulation

Summary The three-dimensional structure of the rat thymus was studied by combined scanning and transmission electron microscopy. The thymus consists mainly of four types of cells: epithelial cells, lymphocytes, macrophages, and interdigitating cells (IDCs).The epithelial cells form a meshwork in the thymus parenchyma. Cortical epithelial cells are stellate in shape, while the medullary cells comprise two types: stellate and large vacuolated elements. A continuous single layer of epithelial cells separates the parenchyma from connective tissue formations of the capsule, septa and vessels. Surrounding the blood vessels, this epithelial sheath is continuous in the cortex, while it is partly interrupted in the medulla, suggesting that the blood-thymus barrier might function more completely in the cortex.Cortical lymphocytes are round and vary in size, whereas medullary lymphocytes are mainly small, although they vary considerably in surface morphology.Two types of large wandering cells, macrophages and IDCs, could be distinguished, as well as intermediate forms. IDCs sometimes embraced or contacted lymphocytes, suggesting their role in the differentiation of the latter cells.Perivascular channels were present around venules and some arterioles in the cortico-medullary region and in the medulla. A few lymphatic vessels were present in extended perivascular spaces.The present study suggests the possible existence of two routes of passage of lymphocytes into the general circulation. One is via the lymphatics, while the other is through the postcapillary venules into the blood circulation. Our SEM images give evidence that lymphocytes use an intracellular route, i.e., the endothelium of venules.     

Structure of sinuses in the human lymph node

Summary A casting technique has been employed to display in three dimensions, the lymphatic microcirculation within the human lymph node. The casting compound filled the marginal sinus, and diffusely permeated the cortical lymphoid parenchyma. However, deep within the lymph node in the medullary region, the medium remained within the limits of the sinus walls. The casts showed well-defined channels appearing similar to vessels. These converged into larger vessels, which drained into efferent lymphatics leaving the node at the hilus.Electron microscopic examination showed that the outer wall of the marginal sinus and the trabecular side of trabecular sinuses had an intact, continuous endothelium with a basement membrane. However, gaps were present in the inner wall of the marginal sinus, as well as in the parenchymal wall of the trabecular sinus. In the medulla, the sinuses were lined by endothelial cells which appeared similar to macrophages. The sinus lining was incomplete and possessed numerous perforations. These observations indicated that sinus walls adjacent to connective tissue served as a barrier to cell movement, but those adjacent to a large lymphoid cell population had gaps, with cells in apparent transit between sinus lumen and parenchyma.     

Anatomy of the renal pelvis in the hamster.

The hamster renal pelvis has been studied by means of low-power light microscopy, scanning electron microscopy and morphometric analyses. The results of this study are highly suggestive that the contact of pelvic urine with the other medulla as well as with the inner medulla may be an important aspect of final urine formation. The outer medulla constituted nearly 50% of the total pelvic surface area, with the inner stripe of the outer medulla more than twice the pelvic surface area of the outer stripe of the outer medulla. The large outer medullary pelvic surface area was accounted for by the elaboration of the upper pelvic walls into peripelvic columns, opercula ("secondary pyramids"), fornices and secondary pouches. A thin simple-squamous to low cuboidal pelvic epithelium separated pelvic urine from outer medullary parenchyma. The inner medulla which constituted about one quarter of the total pelvic surface area was covered by a cuboidal to columnar pelvic epithelium which appeared morphologically similar to the papillary collecting duct epithelium. Tubules and capillaries of the inner medulla did not appear as closely juxtaposed to the pelvic epithelium as did those of the outer medulla. Cortical tissue comprised only 11.7% of the total pelvic surface area and was covered by transitional epithelium similar to that of ureter and bladder. The previously reported impermeability of this epithelium suggests that pelvic urine contact with the cortex is unimportant in final urine formation. The rich layer of smooth muscle under the transitional epithelium probably functions to move urine into and out of the pelvis during pelvic peristalsis, which has been observed in vivo.     

Renal morphology of the hook-lipped African rhinoceros, Diceros bicornis, Linnaeus

The kidney of Diceros bicornis has about 60 lobes, all appearing peripherally. These are separated by interlobar septa, except for small septal defects through which tubules pass. Renal capsule and interlobar septa are fibromuscular and contain small blood vessels. The kidney is about 65% cortex. It contains about 12.5 x 10(6) glomeruli, which form about 7% of the cortical mass and 4.6% of the renal mass. Diameter of a glomerular capsule is about 244 microns, there being no difference in size across the cortex in these adults. The ureter bifurcates into a cephalic and a caudal, fibromuscular, urothelial-lined conduit, into which open about 23 urothelial-lined infundibula. The common large collecting duct, or tubus maximus, of every lobe opens at the apex of its infundibulum. Two tubi may join into one infundibulum. The tubi and their terminal collecting ducts (of Bellini) are part of the inner medulla. Musculature of conduits and infundibula is largely longitudinal. The calyx may be represented by a circular muscle bundle near the apex of every infundibulum. The large intralobar veins are partly adherent to their infundibulum and calyx and receive arcuate veins via valved orifices. Most branches of the renal artery enter via the interlobar septa. Within a septum they branch again and also supply numerous perforators, which thence enter the cortex. Remaining branches of the renal artery enter cortex directly from without. A fibromuscular scaffolding lies deep to arcuate veins where they contact medulla. Where these veins contact cortical tubules; however, their walls become merely endothelium, like the walls of the interlobular veins.     

Mechanical properties of the dog renal capsule.

The renal capsule is an important determinant of whole kidney volume/pressure relationships. To gain further insights into its possible role we examined the mechanical properties of the dog renal capsule using standard materials testing procedures. From each of four locations on the kidney surface, the following mechanical properties of the renal capsule were determined: elastic modulus (force/unit of cross-sectional area theoretically required to double the length of the specimen), tensile stiffness (force/unit width theoretically required to double the length of the specimen), ultimate strength (stress at time of fracture of the specimen), and maximum strain (percent strain at time of the fracture of the specimen). We found that the elastic modulus of the renal capsule from all capsular sites was substantially greater than values previously reported for dog aorta. The stiffness of the capsule covering the anterior-posterior surface of the kidney was found to be about 50% greater than the stiffness of the capsule covering the lateral and polar surfaces of the kidney. The ultimate strength of the anterior-posterior capsule was significantly greater than that of the lateral or polar capsule. This finding may explain the clinical observation that the spontaneous rupture of the renal capsule and parenchyma associated with the acute swelling of transplant rejection is confined almost exclusively to the lateral and polar portions of the renal capsule and cortex. The mean maximum strain at each capsular site was about 35%. This degree of circumferential expansion corresponds to about a doubling of kidney volume. Thus, this observation suggests that the renal capsule is at risk to undergo spontaneous rupture when renal volume increases of this magnitude are observed.     

The origin of the dendritic reticulum cell

Summary ¹⁴C-5,6-DHT-Melanin was injected into the left lateral ventricle of adult rats and its fate followed by light and EM autoradiography and by TEM of structures identified as labeled in preceding light micrographs. Shortly after injection, melanin particles were seen ingested by supraependymal and epiplexus cells, by cells residing in the pia-arachnoid, i.e. free subarachnoidal cells and perivascular cells, and by subependymally located microglia-like cells with intraventricular processes. Up to day four, an increase in the number of labelled phagocytes in the CSF was noted which transformed into typical reactive macrophages. After this time, many intraventricular melanin-laden phagocytes formed rounded clusters; cells of such clusters were subsequently found to invade the brain parenchyma by penetrating the ependymal lining and to accumulate in the perivascular space of brain vessels. ¹⁴C-Melanin-storing macrophages were found in the marginal sinus of the deep jugular lymph nodes suggesting emigration of CNS-derived phagocytes via lymphatics or prelymphatics that contact the subarachnoidal space compartment. This does not exclude the possibility that some of the macrophages leave the brain via the systemic circulation by penetrating the vascular endothelium; these may be disposed of in peripheral organs other than the lymph nodes.The ability of supraependymal, epiplexus, free subarachnoidal and perivascular cells in the pia and of subependymal microglia cells to accumulate synthetic melanin by phagocytosis suggests that these cells are local variants of the same type of resting potential phagocytes of the mammalian brain. The present study shows that ¹⁴C-5,6-DHT-melanin is an ideal phagocytic stimulant and marker for phagocytosis.Supported by grants from the Deutsche Forschungsgemeinschaft     

MMP-independent role of TIMP-1 at the blood brain barrier during viral encephalomyelitis

Infection of the CNS (central nervous system) with a sublethal neurotropic coronavirus (JHMV) induces a vigorous inflammatory response. CD4⁺ and CD8⁺ T cells are essential to control infectious virus but at the cost of tissue damage. An enigma in understanding the contribution of T cell subsets in pathogenesis resides in their distinct migration pattern across the BBB (blood brain barrier). CD4⁺ T cells transiently accumulate within the perivascular space, whereas CD8⁺ T cells migrate directly into the CNS parenchyma. As MMPs (matrix metalloproteinases) facilitate migration across the glia limitans, specific expression of the TIMP (tissue inhibitor of MMPs)-1 by CD4⁺ T cells present in the perivascular cuffs suggested that TIMP-1 is responsible for stalling CD4⁺ T cell migration into the CNS parenchyma. Using TIMP-1 deficient mice, the present data demonstrate an increase rather than a decrease in CD4⁺ T cell accumulation within the perivascular space during JHMV infection. Whereas virus control was not affected by perivascular retention of CD4⁺ T cells, disease severity was decreased and associated with reduced IFNγ (interferon γ) production. Moreover, decreased CD4⁺ T cell recruitment into the CNS parenchyma of TIMP-1 deficient mice was not associated with impaired T cell recruiting chemokines or MMP expression, and no compensation by other TIMP molecules was identified. These data suggest an MMP-independent role of TIMP-1 in regulating CD4⁺ T cell access into the CNS parenchyma during acute JHMV encephalitis.     

The stroma of the thymus of the rat: morphology and antigen diffusion, a reconsideration

This work reconsiders aspects of the morphology of the capsule, of the blood vasculature, of the distribution of reticular fibers, and of the diffusion of intramediastinally injected antigens in the stroma of the thymus of the rat. This was done by an analysis of standard sections of normal thymuses, of sections of thymuses perfused with colloidal carbon, of silver-impregnated sections, and of sections of thymuses of rats injected intramediastinally with a fluorescent antigen or intravenously with Trypan blue, and by electron microscopy of the thymic capsule. The capsule consisted of two layers: an outer layer covering the entire periphery of a thymic lobe, and an inner layer which outlined the entire convoluted peripheral cortex of a lobe. Cortical vessels entered the capsule and septa in which they formed a capillary network. These capsular capillaries were fenestrated and leukocytes were often present near them. Adipocytes were also seen near these vessels in some areas of the capsule, and often at the bases of septa and trabeculae. Furthermore, much of the medulla had a dense network of coarse reticular fibers, whereas the remainder of the medulla and the cortex contained a loose network of fine fibers stretching out from the capsule, septa, and trabeculae. Intramediastinally injected fluorescent antigens were observed to spread in the capsule and septa and to diffuse in the fiber networks stretched across the cortex and the medulla. Fluorescence also highlighted cortical reticular cells but not the thymocytes. Intravenously injected Trypan blue stained the capsule, the septa, the cortical reticular cells, and the autofluorescent cells outlining the corticomedullary junction of each lobule. The unusual penetration of capillaries from the thymic parenchyma into the thymic capsule suggested that the capsular capillaries participate in peculiar thymic events, such as the recruitment of blood stem-cells. It is concluded that small amounts of blood antigens normally exude from capsular capillaries and diffuse into the fibers extending from the capsule across the cortex. The phenomenon would be increased under conditions causing thymic involution. An explanation is proposed to account for the development of involution which involves the exudation of antigens from the capsular capillaries. A comparable mechanism could also account for the development of a particular experimental immune tolerance.     

Application of fluorescent dextrans to the brain surface under constant pressure reveals AQP4-independent solute uptake

Extracellular solutes in the central nervous system are exchanged between the interstitial fluid, the perivascular compartment, and the cerebrospinal fluid (CSF). The “glymphatic” mechanism proposes that the astrocyte water channel aquaporin-4 (AQP4) is a major determinant of solute transport between the CSF and the interstitial space; however, this is controversial in part because of wide variance in experimental data on interstitial uptake of cisternally injected solutes. Here, we investigated the determinants of solute uptake in brain parenchyma following cisternal injection and reexamined the role of AQP4 using a novel constant-pressure method. In mice, increased cisternal injection rate, which modestly increased intracranial pressure, remarkably increased solute dispersion in the subarachnoid space and uptake in the cortical perivascular compartment. To investigate the role of AQP4 in the absence of confounding variations in pressure and CSF solute concentration over time and space, solutes were applied directly onto the brain surface after durotomy under constant external pressure. Pressure elevation increased solute penetration into the perivascular compartment but had little effect on parenchymal solute uptake. Solute penetration and uptake did not differ significantly between wild-type and AQP4 knockout mice. Our results offer an explanation for the variability in cisternal injection studies and indicate AQP4-independent solute transfer from the CSF to the interstitial space in mouse brain.     

The fibromuscular basket (Sporta perimedullaris musculosa) in renculi of kidneys from the ringed seal, Phoca hispida (Pinnipedia)

The medulla of renculi from kidneys of Ringed seals (Phoca hispida) is completely enclosed by cortex except at the hilum. Within the renculus, the fibromuscular coat of the calyx separates from the transitional epithelium at the level of the corticomedullary junction, where the intrarencular arteries also diverge into the parenchyma. Flat ribbons of this stromal tissue form an arborized framework near the medullary side of the intrarencular arteries and the larger of the arcuate arteries derived from them. The ribbons, which are clearly distinct from periarterial connective tissue, are composed of coarse collagenous fibers, elastic fibers, and smooth muscle cells, all oriented in the direction of the long axes of the ribbons, and myofibroblasts. The proportion of smooth muscle cells decreases and that of myofibroblasts increases with increasing distance from the calyx. At the base of the medullary pyramid, the elements of the framework diminish in width and ultimately blend with the surrounding interstitial tissue. The stromal framework, or basket, is homologous with the Sporta perimedullaris musculosa of cetacean kidneys.     

Drainage of macromolecules from the Caudato-Putamen of rat brain

To study the drainage of interstitial fluid and macromolecules from the brain parenchyma, an improved method was developed to inject tracers including Chinese ink in group I and phycoerythrin (PE) in group II into the right caudato-putamen of rat brain. Rats were sacrificed on the 1st, 3rd, 7th, 14th, 21st day after injection in group I and at the 0.5, 1, 2, 5, 24 hour in group II. Distribution of tracers was observed by electron microscopy and fluorescence confocal microscopy. The results showed that tracers distributed diffusely in the white matter at all time points whereas they spread selectively along perivascular spaces in the gray matter by 7 days (d) in group I and 5 hours (h) in group II. Chinese ink was ingested by perivascular phagocytes by 7 d after ink injection. The endothelial cells of capillaries in the gray matter had fluorescence staining in cytoplasm and no staining in nuclei by 24 h after PE injection. Animals in group II were stained with tracers in lateral ventricles, bilateral cervical lymph nodes, and the wall of carotid arteries. These results demonstrated that [1] the macromolecules could be cleared from the caudato-putamen through extracellular space of the neuropil in the white matter and perivascular space in the gray matter, [2] perivascular phagocytes and endothelial cells of capillaries played important roles in clearing macromolecules from the perivascular space, and [3] cervical lymph nodes were involved in draining macromolecules from the brain parenchyma.     

Blood vessels and excretory apparatus of the kidney in some wild animals.

On 110 preparations of the kidney in some wild animals (hare, fox, wolf, bear, boar and chamois), the blood vessels and the excretory apparatus were studied by dissection, injection-corrosion and microscope. Only the bear has a markedly split kidney, whereas the kidneys of the other animals are unsplit. In the fox there is an obvious split of the renal artery into anterior and posterior branches which supply the anterior and posterior portion, respectively, of the renal parenchyma, being separated from each other, so that we may speak of an anterior and posterior kidney. In the fox, wolf, hare and chamois the interlobar arteries pass through the renal calices in a loop composed of adipose tissue, invested by the epithelium of the renal calyx. The renculi of the bear kidney show complete autonomy in relation to the blood vessels as well as in relation to the excretory apparatus. The relation of the surface of the excretory apparatus to the whole kidney was studied. Thus we have found the fox to have relatively the largest excretory apparatus, whose surface amounts to 31% of the whole kidney. In the remainder of the animals investigated this percentage is considerably less, ranging from 21.7% (boar) to 26.7% (bear).     

CXCL12 limits inflammation by localizing mononuclear infiltrates to the perivascular space during experimental autoimmune encephalomyelitis

The inflammatory response in the CNS begins with the movement of leukocytes across the blood-brain barrier in a multistep process that requires cells to pass through a perivascular space before entering the parenchyma. The molecular mechanisms that orchestrate this movement are not known. The chemokine CXCL12 is highly expressed throughout the CNS by microendothelial cells under normal conditions, suggesting it might play a role maintaining the blood-brain barrier. We tested this hypothesis in the setting of experimental autoimmune encephalomyelitis (EAE) by using AMD3100, a specific antagonist of the CXCL12 receptor CXCR4. We demonstrate that the loss of CXCR4 activation enhances the migration of infiltrating leukocytes into the CNS parenchyma. CXCL12 is expressed at the basolateral surface of CNS endothelial cells in normal spinal cord and at the onset of EAE. This polarity is lost in vessels associated with an extensive parenchymal invasion of mononuclear cells during the peak of disease. Inhibition of CXCR4 activation during the induction of EAE leads to loss of the typical intense perivascular cuffs, which are replaced with widespread white matter infiltration of mononuclear cells, worsening the clinical severity of the disease and increasing inflammation. Taken together, these data suggest a novel anti-inflammatory role for CXCL12 during EAE in that it functions to localize CXCR4-expressing mononuclear cells to the perivascular space, thereby limiting the parenchymal infiltration of autoreactive effector cells.     

Perivascular cells in cartilage canals of the developing mouse epiphysis

Morphological variability among perivascular cells adjacent to cartilage matrix during the elongation of canals through both uncalcified and calcified matrix has not been reported. Cartilage canals were located in distal femoral epiphyses of 5- to 7-day-old mice and identified as vascular channels arising from perichondrial surfaces along the condyles and intercondylar fossae. Three stages of canal development were identified based on the length of canals and on characteristics of chondrocytes and matrix surrounding the canals. Superficial canals terminated in uncalcified matrix of resting cartilage; intermediate canals terminated in matrix containing hypertrophic chondrocytes; deep canals terminated in calcified matrix. The ultrastructural morphology of perivascular cells in contact with the matrix varied in the three stages. Cells resembling fibroblasts and vacuolated macrophages were present adjacent to the uncalcified matrix in superficial canals. At the tips of intermediate canals, cells resembling fibroblasts were larger, contained numerous lysosomes and phagolysosomes, and were in intimate contact with the matrix. At the tips of deep canals, chondroclasts with ruffled borders and clear zones contacted the calcified matrix. The results indicate that 1) mouse epiphyses provide a suitable model for studying cartilage-canal perivascular cells, 2) calcification of cartilage matrix occurs along the course of the canal, and 3) the morphology of perivascular cells in contact with the matrix may be determined, in part, by matrix calcification.     

Structure and ultrastructure of leaf and calyx glands in Galphimia brasiliensis (Malpighiaceae)

The present study describes the anatomical structure of calyx and leaf glands in Galphimia brasiliensis and analyzes the mechanism of secretion. The glands are marginal and suprabasal, cup-shaped, sessile, and scarcely visible with the naked eye. Light microscopy reveals the following features: a thin, smooth cuticle; unistratified secretory cells; subglandular parenchyma; and vascular bundle supply composed of phloem and xylem with abundant druses of calcium oxalate. Transmission electron microscopy reveals the presence of secretory cells with conspicuous nuclei, dense cytoplasm, lipid droplets, numerous vesicles, mitochondria, Golgi, rough endoplasmic reticulum (RER), and elongated plastids with osmiophilic contents. The secretion reaches the apoplastic space and accumulates beneath the cuticle. Finally, the viscous, translucent exudate is eliminated by mechanical rupture of the cuticle. Histochemical analysis confirms that lipids are the main constituent. Small amounts of polysaccharides were also identified.     

Neural networks in the mushroom bodies of the honeybee

The Kenyon cells (K cells) or intrinsic neurones of the honeybee's mushroom bodies are organised as a series of arrays. In the calyces the arrays form concentric rings that are represented by rectilinear layers in the α and β lobes. The inputs to the calyces have been revealed by intraneuropilar cobalt injection into the optic and antennal lobes. Neurones from the medulla project to the collar neuropil of the calyx while the relay neurones of the antennal lobe project to the lip neuropil of the calyx. Extrinsic neurones of unknown polarity penetrating the α and β lobes have branching patterns that reflect the layered pattern of the intrinsic neurones. The study illustrates the feasibility of producing a fine grain map of the optic lobe and antennal lobe inputs to the mushroom bodies. It is suggested that the map could be produced by making cobalt injections into individual identified antennal glomeruli and at known sites in the medulla retinotopic mosaic.