Lympho-granulocytic tissue associated with the wall of the spiral valve in the African lungfish Protopterus annectens

We describe the structure of the lympho-granulocytic tissue associated with the wall of the spiral valve of the African lungfish Protopterus annectens. The study was performed under freshwater conditions and after 6 months of aestivation. The lympho-granulocytic tissue consists of nodes surrounded by reticular tissue. The nodes are formed by an outer and an inner component separated by a thin collagenous layer. The outer component is a reticular-like tissue that contains two types of granulocytes, developing and mature plasma cells and melanomacrophage centres (MMCs). The inner component, the parenchyma, contains a meshwork of trabeculae and vascular sinusoids and shows dark and pale areas. The dark areas contain diffuse lymphoid tissue, with a large number of mitoses and plasma cell clusters. The pale areas contain a small number of macrophages and lymphocytes. Macrophages and sinus endothelial cells are filled with haemosiderin granules and appear to form part of the reticuloendothelial system of the lungfish. The reticular tissue houses granulocytes, plasma cells and MMCs and might serve for the housing and maturation of cells of the white series. After aestivation, the nodes undergo lymphocyte depletion, the suppression of mitosis, granulocyte invasion and the occurrence of cell death. By contrast, few histological changes occur in the reticular tissue. Whereas the nodes appear to be involved in lymphocyte proliferation and plasma cell maturation, the function of the reticular tissue remains obscure.     

Post-hatching development of the thymic epithelial cells in the rainbow trout Salmo gairdneri: an ultrastructural study

This study reports the ultrastructure of subpopulations of epithelial cells of the thymic parenchyma during the post-hatching development of the rainbow trout, Salmo gairdner, kept at 14 degrees C. At hatching, the thymus contained a small number of medium and large thymocytes interspersed among three different types of epithelial cells: (1) epithelial cells adjacent to the connective tissue capsule; (2) ramified dark epithelial cells with electron-dense cytoplasm; and (3) pale electron-lucent epithelial cells displaying secretory-like features. All these cells types were anchored to one another by desmosomes and had apparently differentiated from the pharyngeal epithelium. At 4 days after hatching, the thymus enlarged, and numerous gaps occurred between the cell processes of contiguous epithelial cells adjacent to the capsular connective tissue. In 21-day-old trout, thymic trabeculae developed carrying blood vessels, and a subcapsular zone became evident containing lymphoblasts and large subcapsular epithelial cells. In 30-day-old trout, an outer thymic zone developed consisting of spindle-shaped epithelial cells which formed a dense network. At this stage, scattered cystic cells, which apparently differentiated from the pale epithelial cells, were present.     

Macrophages and epithelial cells of the thymus gland. An ultrastructural study in the natterjack, Bufo calamita

In the present study, the ultrastructure of the stromal components, basically epithelial elements and macrophages, of the thymus of adult natterjacks, Bufo calamita has been analyzed. A network of stellate epithelial-reticular cells joined together by desmosomes, constitutes the main component of the thymic parenchyma in both cortex and medulla. In the medulla pale, electron-lucent epithelial cells, sometimes showing surface interdigitations, are striking elements. Moreover, uni- and multicellular epithelial cysts appear in the thymic medulla as well as granulated cells of possible endocrine significance. Remarkably, isolated or grouped gland cells whose morphology and cytoplasmic content resemble that of the skin glands, were occasionally found. Finally, macrophages, multinucleated giant cells and dendritic-like cells, the latter intimately associated to lymphocytes, occur in the thymus of Bufo calamita. The most remarkable morphologic characteristics of all those non-lymphoid cell types, as well as their possible functional significance are comparatively discussed with available information on the amphibian and higher vertebrate thymic cytoarchitecture.     

The ecdysial gland of the spider crab,Libinia emarginata (L). I. Ultrastructure of the gland in the male

The ecdysial glands of mature male Libinia emarginata are pale, yellowish organs composed of lobes of epithelial cells having oval nuclei which are often eccentric and which have one or two nucleoli containing amorphous granular material and coarse strands. The plasma membrane bordering the basal lamina consists of invaginations containing microtubules which may serve to increase the surface area for metabolic exchange. Masses of smooth endoplasmic reticulum and associated vesicles are scattered throughout the cytoplasm. Two or more vacuoles may coalesce. Larger vesicles lie close to the cell surface. Numerous mitochondria with tubular cristae surround the nucleus and frequently are associated with SER. A few Golgi complexes consisting of flattened sacs, cisternae or vesicles, lipid droplets and free ribosomes were seen. Adjacent plasma membranes may be in close apposition or separated by a space filled with vesicles, granules, or blood or supporting cells. This type of ultrastructure is associated with steroid-secreting cells.     

Pale and dark dichromatism related to microhabitats in a herbivorous tanganyikan cichlid fish,Telmatochromis temporalis

Dichromatism and habitat utilization of a small herbivorous cichlid fish,Telmatochromis temporalis, were studied in a littoral rocky area of Lake Tanganyika. Individuals of both sexes had either pale or dark body coloration that did not change for long periods. Both sexes defended territories from consexuals, and a male had a harem of several females. Pale fish had territories in exposed areas and dark fish in shaded ones. Larger fish were dominant over smaller consexuals and occupied well-illuminated areas. When pale and dark fish were kept in areas with shaded and lighted backgrounds, respectively, they changed body colour within a few weeks. When pale fish were in well-illuminated areas and dark ones in shaded areas, they were cryptic. BecauseTelmatochromis temporalis are very vulnerable prey in coastal areas, dichromatism may function as antipredator camouflage.     

Characterization of cell- and region-specific abnormalities in the epididymis of cathepsin A deficient mice

Cathepsin A (PPCA), a lysosomal carboxypeptidase that functions as a protective protein for alpha-neuraminidase and beta-galactosidase in a multi-enzyme complex, has been shown to be expressed in the epithelial cells of the epididymis. In the present study, the epididymis of PPCA-/- mice from 2 to 10 months of age was compared with those of their wild-type counterparts. Major accumulations of pale vacuoles, corresponding to lysosomes, were noted in principal and narrow/apical cells in PPCA-/- mice, and clear cells also appearing highly vacuolated, were grossly enlarged in size. This was especially evident in the caput and corpus regions, where quantitative analyses confirmed that the epithelium of the tubules in these regions was expanding in profile area. In addition, the base of the epithelium in these regions was often greatly vacuolated, corresponding to cells that presented no identifiable features and appeared to be degenerating. Halo cells dispersed at various levels in the epithelium also appeared to be abnormal, accumulating pale lysosomes. Furthermore, numerous macrophages were observed in the intertubular space of the entire duct, presenting a large size and plethora of pale lysosomes. Taken together, the present data indicate major lysosomal abnormalities in the epididymis of PPCA-/- mice in a cell type and region specific manner. In addition, it is suggested that the compromised halo cells, due to PPCA deficiency within their lysosomes, cannot function properly and as a result there is a recruitment of macrophages in the intertubular space.     

Cytochemical Observations on Malic Dehydrogenase,Lipase, Nonspecific Esterase and Monoamine Oxidase in Chick Liver Cell Cultures Infected with Trichomonas vaginalis*

SYNOPSIS. In trypsin-dispersed chick liver cell cultures malic dehydrogenase activity is localized in granules distributed thruout the cytoplasm of fibroblasts, epithelial cells, and macrophages. A progressive increase of the enzymic activity in all cell culture elements, except for phagocytes whose external or internal surfaces remain in direct contact with the parasites, accompanies infection of the cultures with a relatively pathogenic strain of Trichomonas vaginalis. In such phagocytes most staining for malic dehydrogenase is lost. Epithelial cells and parasite-free macrophages in experimental cultures also have a diffuse cytoplasmic reaction. No lipase activity is present in fibroblasts, but epithelial cells and macrophages in chick liver cell cultures contain numerous reactive granules. A strong diffuse cytoplasmic reaction is found in the epithelial cells and a weaker one in the control phagocytes. In cultures infected with T. vaginalis the enzyme is lost progressively from the epithelium and from those macrophages which have engulfed the parasites or whose external surfaces are invested with the flagellates; however, no significant changes in lipase activity can be found in parasite-free experimental phagocytes. In all cell types found in chick liver cultures, the reaction for nonspecific esterase is localized in cytoplasmic inclusions of varying size, some of which tend to accumulate along the cell membranes and around the nuclei. In addition, a weak diffuse cytoplasmic reaction is seen in the epithelial cells. Most cells in cultures infected with T. vaginalis have a significant increase in esterase activity, the exception being the macrophages which contain parasites within their cytoplasm or those with flagellates applied closely to their external surfaces. Only a few specifically stained granules are retained by such phagocytes. Monoamine oxidase activity is limited to fine granules dispersed in the cytoplasm of fibroblasts, epithelial cells, and macrophages of control cultures. Infection of chick liver cultures with T. vaginalis results in lowered enzyme activity in non-phagocytic cells as well as in macrophages with engulfed flagellates and in those whose external surface are invested with the parasites. The number of reactive inclusions appears to increase in trichomonad-free phagocytes of experimental cultures.     

Phagicola angrense (Digenea: Heterophyidae) as a cause of enteritis in a raccoon (Procyon lotor)

Numerous Phagicola angrense were associated with enteritis in a single male juvenile raccoon (Procyon lotor) live-trapped on Parramore Island, Virginia (USA). The raccoon was weak, ataxic and had melenic soft feces. The carcass was emaciated, pale and had ascites. Mesenteric vessels appeared prominent and the stomach and the intestines contained fetid bloody material. The small intestinal mucosa contained three locally extensive sites of necrosis. Histopathologically, there were numerous small digeneans both attached to the mucosa and free within the lumen. Digeneans were usually found deep within the crypts where the epithelium was markedly attenuated and devoid of epithelial cells at the point of parasite attachment. In the lamina propria there were areas of acute hemorrhage and infiltration with plasma cells and eosinophils. This appears to be the first record of severe enteritis in the raccoon caused by this digenean.     

Collagenase-3 expression in periapical lesions: an immunohistochemical study

Collagenase-3 (matrix metalloproteinase-13) is a metalloproteinase (MMP) that is associated with bone lesions and exhibits variable expression patterns in odontogenic cysts; it may play a role in regulating focal proliferation and maturation of jaw cyst epithelium. We studied the localization, staining intensity and distribution of collagenase-3 in 13 periapical granulomas with epithelium, 16 periapical granulomas without epithelium and 10 radicular cysts using archived formalin fixed, paraffin embedded tissues. A monoclonal antibody against human collagenase-3 was used to evaluate its expression. Immunohistochemical staining intensities of collagenase-3 in all periapical lesions were (?), 4 (10%); (+), 1 (3%); (++), 22 (56%) and (+++), 12 (31%); differences were not statistically significant. Immunohistochemical distribution of collagenase-3 in epithelial cells was (?), 17 (44%); (+), 17 (44%); (++), 5 (13%); in fibroblasts it was (?), 8 (20%); (+), 23 (59%); (++), 8 (21%); in plasma cells it was (?), 7 (18%); (+), 22 (56%); (++), 10 (26%); in macrophages it was (?), 7 (18%); (+), and 15 (38%); and (++), 17 (44%). Statistically significant differences were found in epithelial cells (p = 0.00) and fibroblasts (p = 0.02), whereas differences were not statistically significant for plasma cells and macrophages. Collagenase-3 may play a role in the conversion of a periapical granuloma with epithelium to radicular cyst. MMP's influence not only epithelial rest cell migration, but also invasion of various stromal cells into granulomatous tissue.     

Trace fossils of the Jurassic,blue Lias,Lyme Regis,Southern England

The Lower Jurassic, Blue Lias Formation at Lyme Regis, Dorset, England is rhythmically bedded with symmetrical rhythms consisting of laminated black shale followed by dark then pale marls (sometimes cemented into pale limestones) and back to laminated black shale. Its trace fossil fauna includes nine ichno‐genera and represents a shallow shelf paleoenviron‐ment. The ichnofauna occurs in three assemblages named after consistently occurring ichnogenera. A Chondrites assemblage (diversity = 1 ichnogenus) and a Chondrites‐Arenicolites assemblage (diversity = 2–3 ichnogenera) are typical of dark marls but may penetrate tops of laminated black shales. A thalassi‐noides‐Chondrites‐Arenicolites assemblage (diversity up to 7 ichnogenera) is typical of pale marls or limestones. Blue Lias limestones are diagenetic: dark laminated limestones and pale bioturbated limestones are cemented equivalents of laminated black shales and pale marls, respectively. However, pale burrow fills in darker sediments and vice versa confirm the rhythms are primary. Rhythms represent redox cycles, fluctuating from anoxic laminated black shales lacking mac‐robenthos or infauna (bioturbation index = 0) to oxic, bioturbated, pale marls or limestones with large mac‐robenthos such as Gryphaea or Plagiostoma and diverse infauna (bioturbation index = 5).     

Bone morphology of weanling rats from dams subjected to fluoride

Epithelial cells and surrounding free cells in the choroid plexus were examined cytochemically using filipin to clarify the distribution pattern of cholesterol within plasma membranes. The apical and basal membranes of the choroid epithelial cell are less susceptible to filipin than the lateral epithelial membrane and plasma membranes of adjacent mesenchymal cells such as macrophages and fibroblasts. Apical and basal domains of the epithelial membranes, which are relatively resistant to action of filipin, appear to have a slightly lower cholesterol content. We suggest that the apical and basal membranes may possess a unique membrane fluidity or stability that differs from that of the lateral epithelial, macrophage or fibroblast membranes.     

Post-dispersal predation on Pinus sylvestris seeds by Fringilla spp: ground substrate affects selection for seed color

Background matching might lower the risk of seeds being eaten by seed predators that search visually. In aviary experiments, we analyzed the selection of diff erent-colored seeds by ground-feeding finches (Fringillacoelebs and F.montifringilla) against four naturally occurring forest soil substrates. The substrates were fresh burn (black), 6-year-old burn (brown), mineral soil (pale yellow) and Pleuroziumschreberi feather moss (green). We used color-sorted seeds of Pinussylvestris, a species with a large natural variation in seed color, ranging from pale yellow to black. Although seeds were scattered on the substrates at a density of only 91 seeds m⁻², birds removed seeds effectively. Both bird species found more pale than dark seeds on the fresh burn substrate. F. montifringilla also recovered more pale than dark seeds on the old burn, and more dark than pale seeds on mineral soil. In moss, the birds found very few seeds compared to the other substrates, and there was no color discrimination. P.sylvestris is frequently regenerating after fire, suggesting that dark seeds would be favored under selection from visually searching predators. Fire-adapted conifers with serotinous cones, e.g., Pinuscontorta ssp. latifolia, which spread their seeds primarily on freshly burnt surfaces, produce uniformly black or dark brown seeds. However, regeneration of the non-serotinous P.sylvestris is often extended for several years after a fire, during which substrate color and structure change. This may have helped to maintain variation in seed color. When regeneration of a plant species occurs on a substrate of uniform color, we believe that selection by visually searching seed predators will result in the evolution of cryptic seed color. Received: 16 August 1996 / Accepted: 30 November 1996     

Ultrastructure of the tongue and anterior process of the sublingual plica in four species of venomous snakes

The general histology and ultrastructure of the tongue and anterior process of the sublingual plica of four Taiwanese venomous snakes, the Chinese cobra (Naja naja atra), banded krait (Bungarus multicinctus), Taiwan habu (Trimeresurus mucrosquamatus), and bamboo snake (Trimeresurus stejnegeri stejnegeri) are described. The tongue fork exhibits a mid-dorsal invagination that broadens gradually toward its base. No mid-ventral invagination is observed. The epithelial cells on both dorsal and ventral aspects of the tongue fork have large and small microfacets, micropores and microvilli. The cell size, distribution pattern of the large microfacets, and the number of small microfacets present on both sides of the fork are essentially the same within a species, but vary among species. The function of these ultrastructures on the cell surface might be for the capture of chemical substances. The large microfacets are raised areas of the cell membrane, each with a pale granule contained within. The chemical nature of the pale granule is not yet known. The small pores surrounding the large microfacets are shallow hollows left after the release of the pale granules from the microfacets. The basic histological pattern of the tongue fork of these species is similar, being composed of a mucosal layer outside and dense musculature inside. No taste buds are discernible. The anterior processes are concave-like expansions of the anteriormost portions of the sublingual plicae. The oblique folds and micropapillae of this organ might be helpful for receiving the chemicals collected on the tongue, when the tongue makes contact with the elevated processes. The elevated processes may penetrate the ducts of Jacobson's organs to effect the final transfer.     

Castration-induced prostate epithelial cell apoptosis results from targeted oxidative stress attack of M1142-macrophages

Prostate development and function are regulated by androgens. Epithelial cell apoptosis in response to androgen deprivation is caspase-9-dependent and peaks at Day 3 after castration. However, isolated epithelial cells survive in the absence of androgens. Znf142 showed an on-off expression pattern in intraepithelial CD68-positive macrophages, with the on-phase at Day 3 after castration. Rats treated with gadolinium chloride to deplete macrophages showed a significant drop in apoptosis, suggesting a causal relationship between macrophages and epithelial cell apoptosis. Intraepithelial M1-polarization was also limited to Day 3, and the inducible nitric oxide synthase (iNOS) knockout mice showed significantly less apoptosis than wild-type controls. The epithelial cells showed focal DNA double-strand breaks (DSB), 8-oxoguanine, and protein tyrosine-nitrosylation, fingerprints of exposure to peroxinitrite. Cultured epithelial cells induced M1-polarization and showed focal DSB and underwent apoptosis. The same phenomena were reproduced in LNCaP cells cocultured with Raw 264.7 macrophages. In conclusion, the M1 ₁₄₂-macrophage (named after Znf142) attack causes activation of the intrinsic apoptosis pathway in epithelial cells after castration.     

Infiltration of inflammatory macrophages and neutrophils and widespread pyroptosis in lung drive influenza lethality in nonhuman primates

Severe influenza kills tens of thousands of individuals each year, yet the mechanisms driving lethality in humans are poorly understood. Here we used a unique translational model of lethal H5N1 influenza in cynomolgus macaques that utilizes inhalation of small-particle virus aerosols to define mechanisms driving lethal disease. RNA sequencing of lung tissue revealed an intense interferon response within two days of infection that resulted in widespread expression of interferon-stimulated genes, including inflammatory cytokines and chemokines. Macaques with lethal disease had rapid and profound loss of alveolar macrophages (AMs) and infiltration of activated CCR2⁺ CX3CR1⁺ interstitial macrophages (IMs) and neutrophils into lungs. Parallel changes of AMs and neutrophils in bronchoalveolar lavage (BAL) correlated with virus load when compared to macaques with mild influenza. Both AMs and IMs in lethal influenza were M1-type inflammatory macrophages which expressed neutrophil chemotactic factors, while neutrophils expressed genes associated with activation and generation of neutrophil extracellular traps (NETs). NETs were prominent in lung and were found in alveolar spaces as well as lung parenchyma. Genes associated with pyroptosis but not apoptosis were increased in lung, and activated inflammatory caspases, IL-1β and cleaved gasdermin D (GSDMD) were present in bronchoalveolar lavage fluid and lung homogenates. Cleaved GSDMD was expressed by lung macrophages and alveolar epithelial cells which were present in large numbers in alveolar spaces, consistent with loss of epithelial integrity. Cleaved GSDMD colocalized with viral NP-expressing cells in alveoli, reflecting pyroptosis of infected cells. These novel findings reveal that a potent interferon and inflammatory cascade in lung associated with infiltration of inflammatory macrophages and neutrophils, elaboration of NETs and cell death by pyroptosis mediates lethal H5N1 influenza in nonhuman primates, and by extension humans. These innate pathways represent promising therapeutic targets to prevent severe influenza and potentially other primary viral pneumonias in humans.     

Intracellular survival and innate immune evasion of Burkholderia cepacia: Improved understanding of quorum sensing-controlled virulence factors,biofilm, and inhibitors

Burkholderia cepacia complex (Bcc) are opportunistic pathogens implicated with nosocomial infections, and high rates of morbidity and mortality, especially in individuals with cystic fibrosis (CF). B. cepacia are naturally resistant to different classes of antibiotics, and can subvert the host innate immune responses by producing quorum sensing (QS) controlled virulence factors and biofilms. It still remains a conundrum as to how exactly the bacterium survives the intracellular environment within the host cells of CF patients and immunocompromised individuals although the bacterium can invade human lung epithelial cells, neutrophils, and murine macrophages. The mechanisms associated with intracellular survival in the airway epithelial cells and the role of QS and virulence factors in B. cepacia infections in cystic fibrosis remain largely unclear. The current review focuses on understanding the role of QS-controlled virulence factors and biofilms, and provides additional impetus to understanding the potentials of QS-inhibitory strategies against B. cepacia.     

Long‐term live‐cell imaging reveals new roles for Salmonella effector proteins SseG and SteA

Salmonella Typhimurium is an intracellular bacterial pathogen that infects both epithelial cells and macrophages. Salmonella effector proteins, which are translocated into the host cell and manipulate host cell components, control the ability to replicate and/or survive in host cells. Due to the complexity and heterogeneity of Salmonella infections, there is growing recognition of the need for single‐cell and live‐cell imaging approaches to identify and characterize the diversity of cellular phenotypes and how they evolve over time. Here, we establish a pipeline for long‐term (17 h) live‐cell imaging of infected cells and subsequent image analysis methods. We apply this pipeline to track bacterial replication within the Salmonella‐containing vacuole in epithelial cells, quantify vacuolar replication versus survival in macrophages and investigate the role of individual effector proteins in mediating these parameters. This approach revealed that dispersed bacteria can coalesce at later stages of infection, that the effector protein SseG influences the propensity for cytosolic hyper‐replication in epithelial cells, and that while SteA only has a subtle effect on vacuolar replication in epithelial cells, it has a profound impact on infection parameters in immunocompetent macrophages, suggesting differential roles for effector proteins in different infection models.     

The identification of thymic nurse cells in vivo and the role of cytoskeletal proteins in thymocyte internalization

Much debate has been generated about the existence of thymic nurse cells within the thymus. Until now, the authenticity of an epithelial cell capable of internalizing developing thymocytes within the thymic cortex has been in question. Here, we use the thymic nurse cell-specific monoclonal antibody, ph91, to define the in vivo location of thymic nurse cells. For the first time, thymic nurse cells enclosing several thymocytes were detected in the subcapsular region of the thymic cortex in a “honeycomb-like” configuration. In vitro studies show the internalization process using digitalized time-lapse microscopy. Internalized thymocytes have also been reported to interact with macrophages within the TNC complex. The cytoplasmic interaction between thymocytes and macrophages was detected using time-lapse microscopy. Using fluorescence microscopy, we show polymerization of actin within macrophages at the contact point with thymocytes, which is indicative of an immunological synapse. Microfilaments and microtubules within TNCs were shown to be associated with thymocyte binding and internalization, but neither interacted with macrophages. Also, we provide data to show that thymocytes are actively involved in the internalization process. These experiments show for the first time the existence of thymic nurse cells within the thymic microenvironment. They provide a visual documentation of thymocyte uptake by thymic nurse cells, and define an interaction between thymocytes and macrophages within the TNC complex.     

TRPV2 expression in rat oral mucosa

The oral mucosa is a highly specialised, stratified epithelium that confers protection from infection and physical, chemical and thermal stimuli. The non-keratinised junctional epithelium surrounds each tooth like a collar and is easily attacked by foreign substances from the oral sulcus. We found that TRPV2, a temperature-gated channel, is highly expressed in junctional epithelial cells, but not in oral sulcular epithelial cells or oral epithelial cells. Dual or triple immunolabelling with immunocompetent cell markers also revealed TRPV2 expression in Langerhans cells and in dendritic cells and macrophages. Electron microscopy disclosed TRPV2 immunoreactivity in the unmyelinated and thinly myelinated axons within the connective tissue underlying the epithelium. TRPV2 labelling was also observed in venule endothelial cells. The electron-dense immunoreaction in junctional epithelial cells, macrophages and neural axons occurred on the plasma membrane, on invaginations of the plasma membrane and in vesicular structures. Because TRPV2 has been shown to respond to temperature, hypotonicity and mechanical stimuli, gingival cells expressing TRPV2 may act as sensor cells, detecting changes in the physical and chemical environment, and may play a role in subsequent defence mechanisms.     

EMMA-4 analysis of iron in cells of the thymic cortex of a weaver-bird (Quelea quelea).

Combined morphological and analytical studies with the EMMA-4 analytical electron microscope have enabled very early erythroid cells to be identified within the cortex of enlarging thymic lobes of Quelea quelea. These early erythroid cells have pale cytoplasm (sometimes with ferritin-like crystals present), slightly pachychromatic nuclei and have fewer cell organelles (mitochondria) than lymphocytes. Counts for iron were approximately 70% lower than counts from mature erythrocytes found free in the cortex. Iron was also recorded from some epithelial reticular cells and pyknotic nuclei; no iron was recorded from small lymphocytes (thymocytes) in the cortex. The presence of very early erythroid cells is a further indication that erythropoiesis occurs in situ in the avian thymus.