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Chapter 4Ultrastructure of Esophagus of Juveniles of
the Soybean Cyst Nematode, Heterodera glycines
Figure Captions
Figure 68. Longitudinal
diagram of esophageal region of a second-stage juvenile of Heterodera
glycines.
Figure 69. Terminology
of body regions of nematodes citing the triradate symmetry of
esophagus. Reprinted with permission of Dr. A. Coomans and The
Société Royale Zoologique de Belgique from Annales
de la Société Royale Zoologique de Belgique 108:115117,
1979.
Figure 70. Diagram
of dorsal gland ampulla valve or end apparatus in closed position.
Note that the infolded valve membrane curves into the valve duct
lumen.
Figure 71. Diagram
of one of two subventral gland ampullae with valve or end apparatus
in an open position.
Figure 72.
Longitudinal section of anterior procorpus region of esophagus
of Heterodera glycines, showing stylet protractor muscles
( Pm) and portions of their cell bodies ( PmC).
Tonofilamentous ( tf) regions of protractor muscle cell
bodies are separated by a delta-shaped, mitochondria-filled ( Mc)
part of a stylet shaft supporting cell ( ShC). N,
nucleus; Sk, stylet knob. Bar=0.5 µm.
Figure 73. In
longitudinal section, stylet shaft cell (ShC) extending
anteriorly and beyond stylet knob (Sk). Pm,
protractor muscle; PmC, cell body of protractor muscle.
Bar=0.5 µm.
Figure 74. In
longitudinal section of two of four secondary muscle (sm)
elements, their cell bodies (smC) are located on either
side of stylet shaft cell body (ShC). EL, esophageal
lumen; Pm, protractor muscle; Sk, stylet knob.
Bar=0.5 µm.
Figure 75. Transverse
section of anterior procorpus just posterior to stylet knobs,
showing cell bodies of stylet protractor muscles (PmC),
stylet shaft tissues (ShT), and secondary muscle elements
(sm). cu, cuticle; EL, esophageal lumen;
N, nucleus; Sm, somatic muscle. Bar=0.5 µm.
Figure 76.
Longitudinal section showing esophageal lumen ( EL), dorsal
esophageal gland ampulla valve ( DAv), stylet knobs ( Sk),
and part of lumen wall ( ELW) with its supporting membranes
( arrows). Lumen wall forms a short branch that ends in
a membranous valve with cuticularized arms (see
fig. 80). Valve membrane is continuous with anterior supporting
membrane of dorsal gland ampulla ( DA). Mt, microtubules.
Bar=0.5 µm.
Figure 77. Medial
longitudinal section of base (VB) of dorsal gland ampulla
valve. Note that microtubules (Mt) of ampulla cytoplasm
end at or near the base of dorsal gland ampulla valve (see
fig. 76). Bar=0.5 µm.
Figure 78. Transverse
section of a distal section of dorsal gland ampulla valve. Bar=0.5
µm.
Figure 79. Transverse
section through base of valve (VB) of dorsal gland ampulla
(DA). Procorpus cells are densely packed with mitochondria
(Mc) and other organelles. EL, esophageal lumen.
Bar=0.5 µm.
Figure 80. Transverse
section of dorsal gland ampulla (DA) showing nearby esophageal
lumen wall (ELW) and complex of cells and support membranes
that are continuous with anterior ampulla membranes. Dorsal surface
of ampulla is apposed to an enlarged neural process (NP)
that contains numerous neurosecretory granules. Cuticularized
support arms (ca) of tetraradiate dorsal gland ampulla
valve (DAv) are shown in closed position. EL,
esophageal lumen. Bar=0.5 µm.
Figure 81.
Longitudinal section of posterior procorpus and anterior metacorpus,
showing dorsal gland extension ( Dx) within procorpus
( prc) and anterior metacorpus ( m). Gland extension
contains irregularly distributed groups of secretion granules
( SG) and dilated, circular-to-elongate, vesicular rough
endoplasmic reticulum ( RER). Narrow anterior region of
metacorpus consists of collar of multidirectional sphincter-muscle
( Spm) elements. Bar=1.0 µm.
Figure 82. Submedian
longitudinal section of junction between procorpus and metacorpus,
showing tangential view of a sphincter-muscle (Spm) complex
with some forward-directed muscle elements (arrows).
Bar=1.0 µm.
Figure 83. Transverse
section of sphincter muscles (Spm) showing lack of direct
contact with esophageal lumen wall (ELW). Bar=1.0 µm.
Figure 84. Transverse
section of anterior region of metacorpus showing a membrane complex
(mc) attached to lumen wall (ELW). Sections
of five of eight nuclei (N) in this plane of metacorpus
are shown among muscle elements (mm) that extend forward
from metacorpus pump wall. Dorsal region of metacorpus is occupied
by dorsal gland extension (Dx) and three neural processes
(NP), one of which is a large neurosecretory cell (NSC).
Bar=1.0 µm.
Figure 85. Cross
section through anterior region of metacorpus, showing esophageal
lumen wall (ELW) surrounded by muscle elements (mm)
of metacorpus pump located slightly posteriad. Dorsal gland extension
(Dx) lies between two metacorpus muscle elements. N,
nucleus; SG, secretion granules. Bar=1.0 µm.
Figure 86.
Longitudinal section through metacorpus showing metacorpus pump
with associated muscles, nuclei, and subventral esophageal gland
ampullae ( SvA) with valves ( SAv). Ventrosublateral
longitudinal section of metacorpus shows outline of a ventrally
directed flange of pump wall ( mP) with associated muscles
( mm) attached to pump chamber wall. Muscles extend outward
to various sectors of basal lamina of metacorpus. Paired ampullae
of subventral esophageal gland extensions are supported by cellular
and membranous complexes ( CMC) just posterior to pump.
NP, neural process; Spm, sphincter muscle. Bar=1.0
µm.
Figure 87. Longitudinal
section through pump valve, showing row of five of ten nerve nuclei
that occur in ventrosublateral sectors of metacorpus. Two arms
of triradiate pump chamber (mP) are shown in longitudinal
sections. N, nucleus; Spm, sphincter muscle.
Bar=1.0 µm.
Figure 88. Enlargement
of fig. 87 metacorpus pump chamber, showing electron-translucent
cuticular walls of esophageal lumen branch (ELB) and
pump arms (PA), which are supported by pair of electron-translucent
cuticular ridges (cuR). Lumen wall extends into subventral
gland ampulla (SvA) and ends as slender electron-translucent
supports in membranous valve (SAv). A cell complex (CMC)
surrounds base of valve. Neurosecretory granules (NG)
are present in cells next to subventral gland ampullae. Some secretion
granules (SG) in an ampulla appear partially empty. Wall
of pump vestibule (PVe) narrows posteriorly to triradiate
closed lumen (TL). In longitudinal section, one arm of
triradiate lumen appears as pair of electron-translucent cuticular
walls separated by electron-opaque lumen. A similar region is
shown in cross section in fig. 96. NP, neural process.
Bar=1.0 µm.
Figure 89. Transverse
section of a closed subventral gland valve in a second-stage juvenile
at a feeding site. Bar=0.5 µm.
Figure 90.
Submedial longitudinal section showing nuclei ( N) of
cells associated with accumulations of neurosecretions lying dorsally
in metacorpus (see fig. 84). Adjacent
dorsal gland extension ( Dx) contains cytoplasm with large
accumulations of rough endoplasmic reticulum ( RER) with
dilated cisternae. Sphincter muscles ( Spm) of anterior
metacorpus are shown in tangential section. mpm, metacorpus
pump muscle; SvA, subventral gland ampulla. Bar=1.0 µm.
Figure 91. Oblique
longitudinal section showing an open subventral gland ampulla
valve (SAv) supported by cuticularized arms and membranes
(M). Contents of open valve are similar to electron-opaque
contents of secretion granules (SG) in ampulla. Nerve
processes (NP) of axons or dendrites surround the ampulla
of subventral gland (SvA). Muscle elements of metacorpus
pump (mpm) extend posteriad, partially surround ampullae,
and terminate near sphincter muscles (Spm). Bar=1.0 µm.
Figure 92.
Transverse section of metacorpus showing triradiate pump chamber
and muscles, nuclei ( N), and somatic or sarcoplasmic
cells. Bar=1.0 µm.
Figure 93. Enlargement
of metacorpus pump of fig. 92, showing attachment of pump muscles
to pump chamber wall. Pair of electron-translucent "ridges"
on each arm of pump wall are shown in figs. 8688 as spinelike
ridges (cuR) that follow contour of spindle-shaped valve.
Paired muscle elements (mpm1, mpm2)
are attached to each other by membrane junctions (MJ)
and to pump walls (mPW) by hemidesmosomes (hd).
Dorsal esophageal gland extension (Dx) has cytoplasmic
components that include secretion granules (SG), dilated
rough endoplasmic reticulum (RER), and microtubules (Mt).
Bar=1.0 µm.
Figure 94.
Transverse section through posterior metacorpus, showing triangular
shape of central esophageal lumen ( EL) and square cross-sectional
shape of its branches ( ELB) that end in each of two subventral
gland ampullae. These cuticular wall structures are connected
with a complex of cells ( CMC) and membrane junctions.
Dx, dorsal gland extension; mpm, metacorpus
pump muscle; Mt, microtubules; NP, nerve process.
Bar=1.0 µm.
Figure 95. Transverse
section showing the merging of the base of a subventral gland
valve with the wall of esophageal lumen. Bar=1.0 µm.
Figure 96. Transverse
section of open valves (SAv) within subventral gland
ampullae slightly posterior to section in fig. 94. Valve walls
have characteristic single and double membrane boundaries (vM)
that confine the electron-opaque material (EOM) which
apparently accumulates from secretory granules (SG).
Cuticularized triradiate esophageal lumen (TL) extends
posteriad from terminus of metacorpus pump vestibule through isthmus
and terminates at esophago-intestinal valve. Triradiate lumen
wall is modified in shape and is stabilized by membrane junctions
and cells associated with dorsal gland extension (Dx)
and subventral gland ampullae (SvA). mpm, metacorpus
pump muscle; NP, nerve process. Bar=1.0 µm.
Figure 97.
One of a series of transverse sections through the isthmus and
the anterior region of esophageal glands. Partially open triradiate
lumen ( TL) shows walls attached to membrane junctions
that extend to sphincter muscles ( Spm). Isthmus is encircled
by cluster of nuclei ( N) that lie in a region between
metacorpus and nerve ring. SvA, subventral gland ampulla.
Bar=1.0 µm.
Figure 98. Anterior
region of nerve ring (NR) lying posteriad to sphincter
muscle (see fig. 103) and surrounded
by cluster of nuclei as shown in fig. 97. Dx, dorsal
gland extension; Svx, subventral gland extension. Bar=1.0
µm.
Figure 99. Central
region of isthmus (i) showing triradiate esophageal lumen
(TL) surrounded by dorsal (Dx) and subventral
gland extensions (Svx). Nerve ring (NR) completely
surrounds isthmus, with more neural processes in dorsal region
than in ventral. Bar=1.0 µm.
Figure 100.
Dorsal esophageal gland (Dg) dominating anterior of gland
lobe, where it is bordered laterally by nuclei and sublaterally
by subventral gland extensions (Svx) that are filled
with prominent secretion granules (SG). VnP,
ventral nerve processes. Bar=1.0 µm.
Figure 101.
Cross section of irregularly shaped anterior glandular lobe (agl).
Lumen is encircled by slender supporting cells and anterior regions
of dorsal (Dg) and subventral glands (Svg).
Bar=1.0 µm.
Figure 102.
Transverse section of gland lobe showing different types of secretion
granules (SG) in the dorsal (Dg) and subventral
glands (Svg). Bar=1.0 µm.
Figure 103.
Oblique longitudinal section through isthmus, showing dorsal gland
extension ( Dx) with nerve ring tissue ( NR) on
either side. Nuclei ( N) adjacent to nerve ring and isthmus
are part of cluster that encircles this region in transverse section
(see figs. 97, 98). Bar=1.0 µm.
Figure 104.
Transverse section showing neurosecretions (NS) within
nerve processes of nerve ring (NR). Electron-opaque sites
(arrows) indicate axon-axon or axon-dendrite synapses.
Secretory granules (SG) synthesized in dorsal gland (Dg)
differ from those in subventral glands (Svg), with the
former being more homogeneous and having less distinct limiting
membranes (M). EL, esophageal lumen. Bar=1.0
µm.
Figure 105.
Longitudinal section of gland lobe showing portions of esophago-intestinal
valve ( Eiv) and secretion granules ( SG) accumulated
in dorsal gland ( Dg) and subventral gland ( Svg).
Bar=1.0 µm.
Figure 106.
Longitudinal section of gland lobe showing terminus of open cuticle-lined
esophageal lumen (EL) and its relationship to intestinal
lumen (IL). Dg, dorsal esophageal gland; DN,
dorsal esophageal gland nucleus; Eiv, esophago-intestinal
valve. Bar=1.0 µm.
Figure 107.
Transverse section slightly anterior to dorsal gland nucleus showing
apposed membranes (arrows) of esophago-intestinal valve
(Eiv). Subventral gland (Svg) cells are slender
and appear like discrete cells that lie adjacent to esophago-intestinal
valve and dorsal esophageal gland (Dg). Bar=1.0 µm.
Figure 108. Transverse
section of central region of dorsal esophageal gland ( Dg)
filled with densely packed, dilated rough endoplasmic reticulum
( RER), mitochondria ( Mc), Golgi apparatus ( GA),
and some secretion granules ( SG). Esophago-intestinal
tissue is composed of three cells ( Eic1, Eic2,
Eic3) and adjoining membrane junctions near
intestinal lumen. The two subventral glands ( Svg1,
Svg2) and the dorsal gland are clearly defined
by their respective cell membranes. Bar=1.0 µm.
Figure 109.
Golgi apparatus (GA) in early stages of formation of
secretion granules, some of which are shown in figs. 108 and 110.
Bar=0.5 µm.
Figure 110.
Transverse section through dorsal esophageal gland (Dg)
and nucleus (DN) with adjacent intestinal lumen (IL)
and intestinal epithelial cells (IEc). SG, secretion
granules. Bar=1.0 µm.
Figure 111. Longitudinal
section of posterior gland lobe, showing nuclei ( N1,
N2) and secretion granules ( SG) of
paired subventral esophageal gland cells ( Svg1,
Svg2). Slender intestinal region ( int)
broadens immediately posterior to gland lobe. IL, intestinal
lumen. Bar=1.0 µm.
Figure 112.
Transverse section of gland lobe showing many membrane-bound granules
(SG) with wide ranges of electron-opacity in the two
subventral gland cells (Svg1, Svg2),
and intercellular membranes (arrows) between dorsal (Dg)
and subventral gland cells. Sm, somatic muscles. Bar=1.0
µm.
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