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Lobster Atlas

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Lobster Atlas

  1. 1. Atlas of Lobster Anatomy and Histology Jeffrey D. Shields, Robert A. Boyd Virginia Institute of Marine Science The College of William & Mary Cite as: Shields, J.D., and R. Boyd, 2014, Atlas of Lobster Anatomy and Histology, Special Papers in Marine Science No. 9, Virginia Institute of Marine Science, The College of William & Mary. [http://hdl.handle.net/10288/21827] Copyright 2014 Special Papers in Marine Science No. 9 1
  2. 2. Production notes This web publication replaces an earlier atlas of lobster anatomy (Famiglietti & Shields 2002). The 2014 atlas has new photomicrographs, labeling and covers additional tissues, as well as an expanded set of photos on the lobster gross anatomy. The older atlas had some issues with the web presentation so we have removed it. The citation for the older version was: Famiglietti, A.P., and J.D. Shields, 2002, An Atlas of Anatomy for the American Lobster, Homarus americanus. www.vims.edu/tcs/Atlas of Lobster Anatomy.pps, The Virginia Institute of Marine Science 2
  3. 3. Preface This is a histological atlas of the most common organs and tissues found in the American lobster, Homarus americanus. The atlas contains photomicrographs from histological sections of healthy tissues. The sections were all cut at 5-6 µm and were all stained with hematoxylin and eosin as in Shields et al. 2012b. The atlas contains pictures of tissues that are readily observed in dissection and several that are commonly affected by diseases. It is not a complete atlas. Several organs are not covered, notably the central nervous system, ventral nerve ganglion, several sensory organs, and organs associated with molting. The atlas may be useful to you for comparisons with other lobster species, or other crustaceans. Three other references offer similar anatomical or histological perspectives for model crustaceans: penaeid shrimp (Bell & Lightner, 1988), the American lobster (Factor, 1995), and the blue crab (Johnson, 1980). Herrick (1895, 1909) undertook histological descriptions of several tissues in the lobster. His classical works are now freely available online. Our lobster atlas arose partially out of the “100 Lobster Project” and partially from an earlier atlas (Famiglietti & Shields 2002). The histological analysis for the “100 Lobster Project” offered an unparalleled perspective on tissues from normal, healthy lobsters and those affected by epizootic shell disease (Shields et al. 2012a,b). This presented an opportunity to revise, expand and rework the earlier atlas into a more comprehensive and accessible pictorial presentation. The current atlas is a work in progress. We foresee possible expansions by adding other tissues, other staining methods, histopathology, and other imaging techniques. If you use the atlas, let us know what could be added or what improvements could be made. In addition, if you find errors, let us know, and we’ll make efforts to fix them. Jeffrey D. Shields Robert A. Boyd Professor of Marine Science Jeff@vims.edu 3
  4. 4. Table of Contents 1. External Anatomy 9. Eye & eyestalk 2. Cuticle 10. Midgut 3. Epidermis 11. Hepatopancreas 4. Heart 12. Male testis 5. Muscle 13. Male posterior vasa deferens 6. Gills 14. Female ovary 7. Hematopoietic tissue Bibliography 8. Antennal gland Acknowledgements 4
  5. 5. Lobster - Dorsal View The exterior gross anatomy of the lobster has been depicted by Hadley (1906), Herrick (1909) and is reviewed in Factor (1995) . 5
  6. 6. Uropods Telson Antennule Abdomen Somite Rostrum Antenna Eye Walking legs Cephalothorax Chela - crusher claw Chela – shredder claw General anatomy using Herrick’s (1895) original artwork 6
  7. 7. Schematic of the internal anatomy from Herrick’s (1909) 7
  8. 8. Lobster - Lateral View 8
  9. 9. Lobster - Anterior View 9
  10. 10. Lateral view of abdomen showing five abdominal segments or somites. som = somite. som Abdomen 10
  11. 11. Swimmerets Swimmerets (pleopods) from abdomen of a male lobster. Table of contents 11
  12. 12. Cuticle Lobster cuticle showing setal follicles and regularly spaced pore canals (arrows). set = setal follicle, bar = 2 mm set set 12
  13. 13. Cross section showing the typical layering of the cuticle (lobster RI079). Note the pore canal running through the cuticle. con = connective tissue, d = epidermis, endo = endocuticle, epi = epicuticle, exo = exocuticle, p = pore canal, bar = 700 µm epi exo endo p d Con Cuticle 13
  14. 14. endo d MB b Basal region of the cuticle showing the endocuticle, membranous layer, and columnar epithelial cells in the epidermis (lobster ME36). B = basement membrane, d = epidermis, endo = endocuticle, MB = membranous layer, bar = 20 µm Cuticle Table of contents 14
  15. 15. Epidermis Dissection of lobster cuticle showing underlying epidermis. d = epidermis, cut = cuticle d cut 15
  16. 16. Lobster epidermis viewed with a dissecting scope. Epidermis shows concentrations of red pigmentation in chromophores. pig = pigmentation, pigC = pigment concentration, bar = () µm. pig pigC Epidermis 16
  17. 17. Epidermal tissue from Lobster ME47. Note the infolding of the premolt epidermis. con = spongy connective tissue, cut = cuticle, d = epidermis, IF = enfolding, RI = reserve inclusion cell, bar = 100 µm. d cut RI IF con Epidermis 17
  18. 18. cut d RI h Detail of the attachment of the tonofibrillae to the cuticle (lobster ME47). The darkly stained tonofibrillae attach the cuticle to the more lightly stained tendinal cells. The tendinal cells attach to muscle (not shown here). con = spongy connective tissue, cut = cuticle, d = epidermis, h = hemocyte, RI = reserve inclusion cell. t = tonofibrillae, tc = tendril cells bar = 20 µm. con t tc Epidermis 18
  19. 19. Epidermal tissue with fixed phagocytes surrounding an arteriole, and large, opaque RI cells (lobster ME43). con = connective tissue, ff = fixed phagocytes, l = lumen of arteriole, n = nerve fiber, RI = reserve inclusion cell, bar = 20 µm. l RI n ff con Epidermis 19
  20. 20. Schematic of the cuticle, epidermis and tegmental gland from Herrick (1895) The cuticle is a living tissue with several means of communicating with the external environment. This schematic shows sensory organs such as setae, pore canals, and other gland canals. 20
  21. 21. Tegmental glands in epidermal tissue, with an oblique section through the smaller tegmental gland (arrow) (lobster 47D). ctg = cell of tegmental gland, du = duct of gland, RI = reserve inclusion cell, TG = tegmental gland, bar = 20 µm. RI du ctg TG Epidermis 21
  22. 22. Detail of tegmental gland duct in epidermal tissue (Lobster ME47). cc = canal cell, con = spongy connective tissue, du = duct of canal cell, L = lumen, RI = Reserve inclusion cell, TGD = tegmental gland duct, bar = 20 µm. TGD RI L con cc Epidermis Table of contents 22
  23. 23. Close up of heart in thoracic coelom taken with a dissecting scope. her = heart tissue. bar = () µm her Heart 23
  24. 24. Low magnification micrograph of a bisected heart showing the pericardium (connective tissue around the heart), heart muscle and hemal sinuses within the heart (Lobster ME24A)., HS = hemal sinus, m = muscle fibers, peri = pericardium, bar = 900 µm peri m HS Heart 24
  25. 25. Pericardium surrounding the heart (lobster ME29A). Fibrous connective tissue cells provide an outer layer of support to the spongey connective tissues of the pericardium. con = connective tissue, fcon = fibrous connective tissue, bar = 50 µm con fcon Heart 25
  26. 26. Transition between pericardium and heart muscle tissue. Note the muscular attachments to the pericardium (lobster ME29A). h = hemocyte, mC = heart muscle fibers, RI = reserve inclusion, TL = transition layer = 50 µm con mC TL RI h Heart 26
  27. 27. Myocardium with extensive reserve inclusion cells (RI), and complex, intercalated myofibrils of cardiac muscle (lobster ME29A). Note the connective fibers between heart muscle fibers. between the longitudinal muscle fibers. cf = connective fibers, myo = heart myofibrils, RI = reserve inclusion cell, bar = 20 µm myo RI cf Heart 27
  28. 28. The intercalated nature of the myofibrils can be seen in the different section planes (longitidunal, cross section, and oblique) shown here (lobster ME29A). cf = connective fibers, myo = myfibrils, bar = 50 µm myo cf Heart myo 28
  29. 29. Nerve tract in heart tissue surrounded by glial cells. Note the muscular tissue from a different plane bisecting the nerve tract (arrow) and the globular material forming in the RI cell. h = hemocyte, myo = myofibrils, gl = glial cells, RI= Reserve inclusion cell. RI-f=Reserve inclusion forming, bar = 20 µm RI RI-f myo gl h Heart Table of contents 29
  30. 30. Muscle Low magnification of skeletal muscle (lobster ME27A). Note the hemal sinuses running longitudinally across the section of skeletal muscle. hs= hemal sinus, ms = skeletal muscle fibers, bar = 200 µm. hs ms 30
  31. 31. ms HS Arteriole in muscular tissue surrounded by fixed phagocytes (lobster ME27A).ff = fixed phagocytes, HS = hemal sinus, L = lumen, ms = skeletal muscle fibers, bar = 20 µm. ff L Muscle Table of contents 31
  32. 32. Gill G Exposed branchial chamber with gill branchiae. G = gill. 32
  33. 33. G Branchial chamber showing epipodites of gill branchiae. Each gill has 3-4 epipodites. Note the attachment of the gill to the base of the pereiopod (arrow). G = gill, br = branchia. br Gill 33
  34. 34. Vertical Section of Gill Low magnification of cross section through the gill (lobster ME29). ch = efferent or afferent hemolymph channel, lam = lamella, bar = 600 µm. lam ch Gill ch 34
  35. 35. st Low magnification longitudinal section through the gill (lobster ME29). Note the trichobranchiate structure of the lamellae on the branchia. Upper lamellae are longitudinally sectioned while the lower lamellae are cross sectioned. Lam=Lamella, st = branchial stem, bar = 200 (10x) Lam Gill Lam 35
  36. 36. Longitudinal section through a gill branchia showing the lamellae connecting to the branchia (Lobster ME29C). Note the hemal sinuses (arrows) below the lamellae. These empty into the afferent and efferent hemolymph channel. Ch=channel of the branchial artery, Con=connective tissue, Lam= gill lamella, bar = 300 µm. Lam Ch Con Gill 36
  37. 37. Lam sin D Detail of lamellae demonstrating their tubular nature. One side of the lamella shows the thin epidermal layer, the other side shows an oblique epidermal layer, with a hemal sinus in the middle. from Lobster ME29C. D = epidermis, Lam = lamella, sin = hemal sinus, bar = 100 µm. Gill D sin 37
  38. 38. hs Longitudinal section of afferent/efferent channel in gill tissue (lobster ME29C). Note the tonofibrillae (arrow) and myofibrils at the base of the the lamella. con = spongy connective tissue, hs = hemal sinus, bar = 50 µm. con Gill 38
  39. 39. Cross section through the lateral portion of lamellae (lobster ME29C). The trabecular cells serve as support cells within the lamellae. cut = cuticle, d = epidermis, h = hemal sinus, trc = trabecular cells, bar = 50 µm. cut d hs trc Gill 39
  40. 40. Cross section of lamella from Lobster ME29C showing thin cuticle and epidermis region. Cut = cuticle, d = epidermis, hs= hemal sinus, trc = trabecular cell, bar = 20 µm. cut d trc hs Gill 40
  41. 41. Branchial region showing effete tegmental gland, and reserve inclusion cells (lobster ME29C). con = connective tissue, ctg = tegmental gland cells, d = epidermis, l = lumen, RI = reserve inclusion cell, bar = 20 µm. l ctg con RI d Gill Table of contents 41
  42. 42. Hematopoietic tissue Exposed thoracic cavity showing pyloric stomach with bands of hematopoietic tissue (arrows). ps = pyloric stomach ps 42
  43. 43. Hematopoietic tissue is the source of hemocytes, or blood cells, in the lobster. The tissue occurs as lobules of cells on the dorsal pyloric stomach, or foregut (Lobster ME24A). Note the nerve tract interspersed between the nodules of hematopoietic tissue. lob = lobule, n = nerve, bar = 100 µm. n lob Hematopoietic tissue n 43
  44. 44. Lobule of hematopoietic tissue surrounded by a rind of accessory columnar epithelial layer (Lobster ME24A). epi = columnar epithelial cells, lob = lobules of tissue with stem cells, bar = 20 µm. lob epi Hematopoietic tissue 44
  45. 45. lob l ff Lobules of hematopoietic tissue with an arteriole running through the tissue (lobster ME24A). ff = fixed phagocytes, l = lumen of arteriole, lob=lobule, bar = 20 µm. Hematopoietic tissue Table of contents 45
  46. 46. Antennal “Green” Gland Antennal gland, or green gland, at the anterior end internally, ventral to the eyes, with a small piece of the bladder attached (arrow). 46
  47. 47. Low magnification of antennal gland showing the organization of the labyrinth tissue and coelomosac (lobster 22A). A small part of the bladder is attached to the labyrinth. The antennal gland functions in excretion and osmoregulation. bl = bladder, ce = coelomosac, ly= labyrinth, bar = 900 µm. ce ly Antennal gland bl 47
  48. 48. Antennal gland at the border between the labyrinth and the coelomosac (lobster 22A). Note the difference in staining coloration between tissues. ce = coelomosac, ly = labyrinth, bar = 300 µm. ly ce Antennal gland 48
  49. 49. d Coelomsac showing squamous epithelial cells arranged in a complex pattern within the gland (lobster ME34A). Note the ducts within the coelomosac and the fibrous connective tissue around the organ. There is no connective tissue within the sac. d = duct, fcon = fibrous connective tissue, le = labyrinth epithelium, bar = 50 µm. fcon le Antennal gland d 49
  50. 50. Border between the labyrinth and coelomosac (lobster ME34A). Note the large duct within the coelomosac (arrow). This could be a connection with the bladder or an arteriole. ce = coelomosac, le = labyrinth, ly = labyrinth, po = podocyte, bar = 100 µm. LY le ce Antennal gland 50
  51. 51. Podocytes in the coelomosac (lobster ME34A). These cells function in excretion and osmoregulation. Note the many brown granules present in the podocyte (arrow). An arteriole is also present. aa = arteriole, hsp = hemal space, h = hemocyte, po = podocyte, bar = 10 µm. po hsp Antennal gland 51
  52. 52. Arteriole in the labyrinth (lobster ME24A). Note the brush border in the lumen of the labyrinth. art = arteriole, bb = brush border, le = labyrinth epithelium, bar = 20 µm. art le Antennal gland bb 52
  53. 53. Labyrinth epithelium (lobster ME34A) with presumptive podocytes (arrows) used in excretion. d = duct, le = labyrinth epithelium, bar = 10 µm. d le Antennal gland 53
  54. 54. ff h le Antennal gland Interdigitation of the secretory labyrinth (lobster ME22A). Sloughed cells are components of the secretory labyrinth. Bb = brush border, h = hemocyte, le = labyrinth epithelium, ss = sloughed epithelial cells, bar = 20 µm. bb h le ss Table of contents 54
  55. 55. Lateral view of lobster eye. Eye and eyestalk 55
  56. 56. Dorsal aspect of lobster eye. Note the cuticle that covers the eyestalk is thicker and has a cuticular pigmentation compared with the thin cuticle that covers the eye proper. eys = eyestalk. eys Eye and eyestalk 56
  57. 57. Eye and eyestalk Gross view of the lobster eyestalk in longitudinal section. The eyestalk was removed, fixed in z-fix, decalcified in formic acid – sodium citrate, then bisected with a razor. Note the increasing thickness of cuticle on the lateral margins of the eye. cut = cuticle, lm = lamina ganglionaris, m = retractor muscle, oma = ommatidia, on = optic nerve region, np = nerve plexus. cut on lm oma m np 57
  58. 58. Low magnification of the lobster eyestalk in longitudinal section (lobster AM 65). The empty space between the cuticle and ommatidia is an artifact of fixation. Note the increasing thickness of cuticle on the lateral margins of the eye. bm = basement membrane of ommatidial region, cut = cuticle, lm = lamina ganglionaris, m = retractor muscle, me = medulla externalis, mi = medulla internalis, mt = medulla terminalis, np = nerve plexus, oma = ommatidia, tg = tegmental glands, bar = 900 µm. cut oma bm lm m tg Eye and eyestalk m on mt mi me np 58
  59. 59. Eye showing ommatidia (bracket) stretching from the cuticle to the basement membrane (lobster RI44). Note the separation of the premolt cuticle from the old cuticle. bm = basement membrane, cc = crystalline cone, cut = cuticle, om = ommatdium, pr = proximal retinula, rh = distal retinula and rhabdom, bar = 300 µm. pr cut cc rh bm Eye and eyestalk om 59
  60. 60. Ommatidia stretching from the cuticle to the basement membrane (lobster RI44). Accessory pigment cells have screening pigments (arrows) around the cones and at the base of the ommatidium. bm = basement membrane, cc = crystalline cone, cut = cuticle, pr = proximal retinula, rh = distal retinula and rhabdom, bar = 100 µm. pr cut cc rh bm Eye and eyestalk 60
  61. 61. Eye and eyestalk Higher magnification of corneal cells (lobster RI53). The open areas are artifacts introduced during processing. ch = corneal hypodermis, cc = crystalline cone, cut = cuticle, bar = 10 µm. cut coc ch Cornea associated with individual ommatidia. Note the screening pigments in the accessory pigment cells (arrows). cr = cornea, cc = crystalline cone, cut = cuticle, om = ommatidia. cut cc om cr 61
  62. 62. Eye and eyestalk -10x Juncture of the ommatidia with the basement membrane in the eye (lobster RI44?). Screening pigments (arrows) surround the proximal rhabdom. Note the vasculature around the basement membrane. bm = basement membrane, hc = vasculature of the hemocoel, pr = proximal retinula, on = optic nerves, bar = 50 µm. pr hc on bm 62
  63. 63. Proximal retinula of the ommatidium (lobster RI44). Note the pigment granules extending above and around the rhabdom, as well as the pigment cell nucleus (arrow). prhc = proximal retinula, rh = rhabdom, bar = 20 µm. rh pr Eye and eyestalk 63
  64. 64. Region of the basement membrane separating the ommatidia from the optic nerve fibers (lobster ME75E). Many pigment granules are present in accessory pigment cells (arrow) that extend into the underlying tissue. bm = basement membrane, g = granules, lc = lacuna of hemocoel, on = optic nerve, bar = 10 µm. bm lc g Eye and eyestalk on on 64
  65. 65. Optic nerve region between the ommatidia and the lamina ganglionaris (lobster AM65). Note the long optic nerve fibers attaching the ommatidia to the lamina ganglionaris and the highly vascular nature of the region. art = arteriole, bm = basement membrane, hs = hemal sinus, lg = lamina ganglionaris, on = optic nerve fibers, bar = 50 µm Eye and eyestalk on art hs lg bm 65
  66. 66. Detail of lamina ganglionaris (lobster RI44). cr = rind of support cells, gl = glial cells, lg = lamina ganglionaris, mx = medulla externalis, onf = optic nerve fiber, bar = 20 µm. onf gl cr mx Eye and eyestalk lg onf 66
  67. 67. Eye and eyestalk Highly vascularized region of the medulla externalis showing optic nerve tracts from the lamina ganglionaris into the region. Specialized rind cells or secretory cells encase the optic nerves as they enter into the medulla internalis (not shown) (lobster AM65). art = arteriole, cr = rind cells, me = medulla externalis, on = optic nerve fibers, bar = 50 µm cr cr art art lg on on me 67
  68. 68. Eye and eyestalk Nerve plexus known as the medulla internalis showing optic nerves entering from the medulla externalis and exiting the medulla internalis (lobster AM65). The cell rind of the medulla internalis is thin and less organized than that of the other medullar ganglia. art = arteriole, cr = rind cells, mi= medulla internalis, on = optic nerve fibers, bar = 100 µm art cr on on mi 68
  69. 69. Eye and eyestalk Nerve plexus known as the medulla terminalis with individual optic nerves entering from the medulla internalis (lobster AM65). Neurosecretory cells (arrows) occur loosely around the outer margin of the medulla internalis and terminalis. The optic nerves continue from here on to the brain of the lobster. art = arteriole, fcon = fibrous connective tissues, cr = rind cells, mt= medulla terminalis, on = optic nerve fibers, bar = 100 µm mt cr on art fcon 69
  70. 70. Eye and eyestalk Dorsal to the medulla terminalis is the sinus gland which controls molting. The lumen of the gland is not shown in this micrograph. Note the numerous neurosecretory cells, the neurilemma and glial cells surrounding the medullae and sinus gland (lobster AM65). gl = fibrous glial cells, mt= medulla terminalis, nsc = neurosecretory cells, sg = sinus gland, bar = 100 µm 70
  71. 71. Dense grouping of tegmental glands near the proximal end of the eyestalk (lobster RI44). Inset shows more detail of a tegmental gland with a common duct. con = connective tissue, cut = cuticle, epi = columnar epithelial cells underlying the cuticle, gran = cuticularized granuloma, tg = tegmental gland, bar = 50 µm. cut con tg gran Eye and eyestalk epi tg Table of contents 71
  72. 72. Gross view of exposed thoracic cavity showing midgut (arrow) running between lobes of the hepatopancreas. Midgut 72
  73. 73. bl ec fcon The food bolus of arthropods is encased in a peritrophic membrane secreted by the midgut. This low magnification view of the midgut shows the epithelium of the organ, the thin layer of supporting tissues around it, and a food bolus with a peritrophic membrane (arrows) around it (lobster RI92). bl = food bolus (gut contents), ec = epithelial cells, fcon = fibrous connective tissue with supporting cells, L = lumen, bar = 1200 µm. L Midgut 73
  74. 74. Midgut The midgut epithelium is columnar with a brush border. Underlying it is a basement layer and a band of reserve inclusion cells interspersed among myofibrils of circular muscle (lobster RI92). bl = food bolus, bm =basement layer, ec = epithelial cell, pm = peritrophic membrane, RI = zone of reserve inclusion cells and myofibrils, bar = 50 µm. bm bl RI ec pm 74
  75. 75. The midgut epithelium is columnar with a brush border. The peritrophic membrane is a thin layer of cuticle that has separated (as an artifact) from the bolus (lobster RI92). bl = food bolus, ec = epithelial cell, L = lumen, pm = peritrophic membrane, bar = 50 µm. ec bl pm L Midgut 75
  76. 76. The midgut has columnar epithelial cells supported on a basement membrane overlying small muscle fibrils and absorptive reserve inclusion cells (lobster RI92). Basal cells (arrows) can be seen among the epithelial cells. Their function is not known. bb = brush border, bm = basement membrane, fcon = fibrous connective tissue, ec = epithelial cells, L = lumen, m = muscle, RI = reserve inclusion cells, bar = 20 µm. ec fcon bm L Midgut bb RI m Table of contents 76
  77. 77. The hepatopancreas is comprised of a mass of individual tubules (arrow) connected through common absorptive ducts to the midgut gland. Hepatopancreas 77
  78. 78. Gross view of hepatopancreas showing individual tubules in longitudinal and cross sections (lobster ME23A). lt = longitudinal section of tubules, xt = cross section of tubule, bar = 600 µm. lt xt Hepatopancreas 78
  79. 79. Hepatopancreatic tissue showing two zones: the E-cell (Embryozellen) zone near the apical end of the tubule, and the digestion zone with B- (Blasenzellen), F- (Fibrillenzellen) and R-cells (Restzellen) near the basal end. bc = B-cell, ec = E-cell, fc = F-cell, fcon = fibrous connective tissue, L = lumen, rc = R-cell, s = B-cell secretion, bar = 100 µm. fcon ec bc L s Hepatopancreas fc rc 79
  80. 80. ec l s Hepatopancreas The stem E-cells in a cross section near the apical end of a hepatopancreatic tubule (lobster ME19). Note the brush border at the apical end of the cells (arrow). bm = basement membrane, ec = E-cell, L = lumen, s = B-cell secretion, bar = 10 µm. bm 80
  81. 81. Hepatopancreatic tubules in cross section (lobster ME19). Note the higher abundance of E-cells in tubules sectioned near the edge of the tissue. This is because the apical portions of these tubules cross the plane of section while tubules within the inner portion of the tissue were sectioned in a more basal region. bc = B-cell, ec = E-cell, L = lumen of tubule, bar = 100 µm. ec bc L Hepatopancreas 81
  82. 82. Secretory B-cells in the hepatopancreatic tubule (lobster ME19). These are holocrine cells in that they secrete their entire contents into the lumen. The nucleus (arrow) is characteristically pushed to the side of the cell. bc = B-cell, L= lumen, s = B-cell secretion, bar = 10 µm. bc s L Hepatopancreas 82
  83. 83. F-cells and R-cells in a hepatopancreatic tubule (lobster ME19). The function of the F-cells is unknown; they may become B-cells. The R-cells function in storage of products . bb = brush border, bm = basement membrane, fc = F cells, L = lumen, rc = R cells, bar = 10 µm. fc rc bm L Hepatopancreas bb 83
  84. 84. Fixed phagocytes surrounding an arteriole (arrow) in the hepatopancreas (lobster ME23A). The fixed phagocytes remove pathogens from the tubules. bm = basement membrane, fc = F-cell, ff = fixed phagocytes, hs = hemal sinus, rc = R-cell, RI = effete reserve inclusion cell, bar = 50 µm. Photo from Shields et al. 2012b. hs rc fc Hepatopancreas rc ff RI bm Table of contents 84
  85. 85. t Thoracic cavity with the heart removed showing testes and hepatopancreas. t = testis, hep = hepatopancreas, m = thoracic muscle hep Male Gonad: Testis m 85
  86. 86. Lobster testis is made up of many lobules (arrow) surrounding a seminiferous duct encased in a thin capsule of fibrous connective tissue cells. Male Gonad: Testis 86
  87. 87. Cross section of testis showing a lobules containing germinative centers in various stages of development (lobster ME23B). A capsule of fibrous connective tissues encases the testis, with thinner connective tissues supporting the lobules internally. l = lobules, f = fibrous capsule, sem = seminiferous duct bar = 300 µm lob Male Gonad: Testis f f sem 87
  88. 88. Testis tissue from lobster ME23B showing non- flagellated mature sperm within a seminiferous epithelium. f = fibrous connective tissues, spg = spermatogonia, spM = mature sperm, s1 = primary spermatocytes, s2 = secondary spermatocytes, sem = seminiferous duct with spermatocytes, bar = 100 µm. f spg s1 s2 spm sem Male Gonad: Testis 88
  89. 89. Non-flagellated mature sperm cells (arrow) in lumen of seminiferous duct of testis (lobster ME23B). spg = Spermatogonia, spM = mature sperm, sem = seminiferous duct wall, bar = 20 µm. sem Male Gonad: Testis 89
  90. 90. Spermatogonia in testis (lobster ME23B). f = fibrous connective tissues, s1 = primary spermatocytes, s2 = secondary spermatocytes, bar = 20 µm. f s1 s2 Male Gonad: Testis 90
  91. 91. Testicular lobules within the testis (lobster ME23B). f = fibrous connective tissues, spg=spermatogonia, sem = seminiferous duct, s1=Primary spermatocytes, s2=Secondary spermatocytes, bar = 100 µm. f sem spg s1 s2 Male Gonad: Testis Table of contents 91
  92. 92. Male Gonad: Ejaculatory Duct Ventral view of lobster showing male gonopore (arrow) and specialized nature of the first pleopod. 92
  93. 93. Close up view of male gonopore (arrow) and male first pleopod. Male Gonad: Ejaculatory Duct 93
  94. 94. Ejaculatory duct dissected from abdomen (circle). Male Gonad: Ejaculatory Duct 94
  95. 95. Low magnification of the ejaculatory duct (lobster ME19). The duct is the posterior-most portion of the posterior vasa deferens. ec = epithelial cells, ms = skeletal muscle, tg = zone of tegmental glands, bar = 1500 µm. ms tg ec Male Gonad: Ejaculatory Duct ec 95
  96. 96. Zone of tegmental glands associated with muscle laterally around the posterior ejaculatory duct (lobster ME19). These glands may provide additional lubrication to the duct or lubrication for molting. fcon = fibrous connective tissue, ms = skeletal muscle, tg = zone of tegmental glands, bar = 50 µm. mS tg fcon Male Gonad: Ejaculatory Duct 96
  97. 97. Cross section of ejaculatory duct containing mature sperm within a waxy spermatophore (lobster ME 12-1). fcon = fibrous connective tissue, RI = reserve inclusion cell, spe = spermatophore, spm = mature sperm, scon = spongy connective tissue, bar = 50 µm. spm ec ri fcon spe Male Gonads: Ejaculatory Duct scon 97
  98. 98. spm Detail of ejaculatory duct with mature sperm in a waxy spermatophore (lobster ME 12-1). ec = columnar epithelial cells, fcon = fibrous connective tissue, RI = reserve inclusion cell, spe = spermatophore, spm = mature sperm, bar = 50 µm. cill ec spe fcon Male Gonads: Ejaculatory Duct 98
  99. 99. Columnar epithelium in the ejaculatory duct (lobster ME19). Note the brush border cill = cilia, ec = epithelial cells, fcon = fibrous connective tissue, mS = skeletal muscle fibers, bar = 20 µm ms fcon ec bb Male Gonads: Ejaculatory Duct Table of contents 99
  100. 100. Female Gonad: Ovary fcon vova ml iml Ovary showing ova in different stages of maturation (lobster ME44B). fcon = fibrous connective tissue ‘rind’, ml = mature lobule of ovary, iml = immature lobule, pre = previtellogenic oocyte, vova = vitellogenic oocyte, bar = 1500 µm. pre 100
  101. 101. Detail of ovarian lobules showing maturing and immature lobules. acb = accessory cell border, fcon = fibrous connective tissue ‘rind’, ml = mature lobule of ovary, iml = immature lobule, pre = previtellogenic ooocyte, vova = vitellogenic oocyte, bar = 300 µm acb pre vova ml iml fcon Female Gonad: Ovary 101
  102. 102. Vitellogenic and previtellogenic oocytes in ovary (lobster ME46B). Note the large yolk (vitellogen) granules forming within the immature oocyte, the numerous granules within the adjacent vitellogenic oocyte, and the chorion (arrows). acb = accessory cell border, n = nucleus of oocyte, nu = nucleolus of oocyte, pre = previtellogenic oocyte, vova = vitellogenic oocyte, , yg = yolk globule bar = 50 µm. acb vova yg pre Female Gonad: Ovary yg n nu 102
  103. 103. Female Gonad: Ovary Detail of accessory cell border between two previtellogenic oocytes (lobster ME46B). acb = accessory cell border, yg= yolk globule, pre = previtellogenic oocyte, bar = 10 µm. acb pre pre yg 103
  104. 104. Yolk globules (vitellogenin) in vitellogenic oocyte (lobster ME46B). Note that some globules contain smaller globules inside them (arrow). yg = yolk globule, bar = 10 µm. yg Female Gonad: Ovary 104
  105. 105. Fibrous tissue around ovary (lobster ME46B). Note the border of accessory cells separating oocytes and the fibrous connective tissue delineating the ovary. acb = accessory cell border, fcon = fibrous connective tissue “rind” around ovary, pre = previtellogenic oocyte, vova = vitellogenic oocyte, yg = yolk globule, bar = 10 µm. acb fcon vova pre yg Female Gonad: Ovary Table of contents 105
  106. 106. Bibliography 106
  107. 107. Bibliography Bell, T. and D. Lightner. (1988) A Handbook of Normal Penaeid Shrimp Histology. World Aquaculture Society, Louisiana. Factor, J. (1995) Biology of the Lobster. Academic Press, New York. Herrick, F.H. (1909) Natural history of the American lobster. Bulletin of the United States Bureau of Fisheries, 29, 149-408 Johnson, P. (1980) Histology of the Blue Crab, Callinectes sapidus. Praeger, New York. Maniscalco, A.M. and J.D. Shields. (2006) Histopathology of idiopathic lesions in the eyes of Homarus americanus from Long Island Sound. Journal of Invertebrate Pathology 91: 88-97 Shields, J.S., K.N. Wheeler, J. Moss. (2012) Histological assessment of lobsters in the “100 Lobster” Project. Journal of Shellfish Research 31: 439-447. Shields, J.D., K.N. Wheeler, J. Moss, B. Somers, K. Castro (2012) The “100 Lobsters” Project: a cooperative demonstration project for health assessments of lobsters from Rhode Island. Journal of Shellfish Research 31: 431-438. Table of contents 107
  108. 108. Acknowledgements We thank Kersten Wheeler and Shelley Sullivan Katsuki for histological processing of lobsters used in the project. Dr. Mike Newman provided a camera for macro photography. Many of the lobster used in the atlas were from the “100 Lobster Project.” This work was supported in part by the National Marine Fisheries Service as the “New England Lobster Research Initiative: Lobster Shell Disease” under NOAA grant NA06NMF4720100 to the University of Rhode Island Fisheries Center, in part by a contract from the Dept. Marine Resources, State of Maine, and in part by a National Science Foundation Research Experience for Undergraduates program grant OCE 0552612 to Drs. L. Schaffner and R. Seitz, VIMS. The views expressed herein are those of the authors and do not necessarily reflect the views of NOAA or any of its sub-agencies. The US Government is authorized to produce and distribute reprints for government purposes, notwithstanding any copyright notation that may appear hereon. Table of contents 108

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