DISTRIBUTION OF DIGESTIVE TUBULES AND FINE STRUCTURE OF DIGESTIVE CELLS OF APLYSIA PUNCTATA (CUVIER, 1803)

The distribution of digestive tubules of Aplysia punctatahas been studied in animals under experimental feeding conditions.Histological analysis of the digestive gland has revealed twotypes of tubules, called tubules A and B. Tubules of type Awere composed of basiphilic cells (calcium, excretory and thincells) and tubules of type B were lined by large digestive cellsand basiphilic cells. The latter occur in small groups, usuallyin the corners of the tubules. Type A tubules are involved inion metabolism and show a diphasic cycle (absorptive and reconstitutive)according to the height and the stage of calcium cells. TypeB tubules are involved in digestive processes and display atetraphasic cycle (holding, absorption, fragmentative and reconstitutive)depending upon the height and the stage of the digestive cells.The tetraphasic cycle was compared with the four categoriesof tubules in bivalves. It is proposed that digestive processesmay be continuous in digestive cells of A. punctata. (Received 16 November 1999; accepted 1 October 2000)     

MONOPHASIC AND DIPHASIC DIGESTIVE CYCLES IN VENERUPIS DECUSSATA AND CHLAMYS VARIA

Feeding and digestive cycles in Chlamys varia and Venerupisdecussata are identified and shown to relate to a tidal cycle.The processes of digestion within the diverticula of the twospecies are very different. The tubules of Venerupis are synchronizedand exhibit a monophasic cycle completed within a twelve-hourperiod, facilitating feeding at the next cycle of the tide.The digestive process of Chlamys requires twenty-four hoursand to accommodate feeding at each twelve-hour tidal cycle thetubules within the diverticula exhibit two different digestivephases simultaneously so that a diaphasic cycle is apparent.The pH of the mantle cavity and regions of the digestive tractand the formation and dissolution of the crystalline style areshown to be related to the tidal cycle. ^*Department of Brewing and Biological Sciences, Heriot-WattUniversity, Chambers Street, Edinburgh, EH1 1HX. (Received 22 March 1978;     

Histological and ultrastructural characterization of the alimentary system of solifuges (Arachnida,Solifugae)

Solifuges are voracious and fast predators. Once having captured a prey item, mostly small arthropods or even small vertebrates, they start feeding on their prey by constant chewing movements with their huge chelicerae. At the same time, they squeeze out the soft tissue that passes the anterior lattice‐like part of the mouthparts. The digestion of the food takes place in the midgut, which is anatomically highly complex. It consists of the midgut tube from which numerous prosomal and opisthosomal diverticula and tubular lateral branches arise. The dimorphic epithelium of the midgut tube and the diverticula is constituted of digestive and secretory cells. The digestive cells are characterized by an apical tubulus system and contain nutritional vacuoles, lipids, spherites, and glycogen. Secretory cells contain a huge amount of rough endoplasmic reticulum and secretory vacuoles. The lateral branches are ultrastructurally similar to Malpighian tubules and are likely involved in excretion. In contrast to the midgut, the epithelium of the hindgut consists of only one type of cell overlain by a thin cuticle. Digested residuals are stored in the hindgut until defecation. J. Morphol., 2010. © 2009 Wiley‐Liss, Inc.     

Cellular growth of host and symbiont in a cnidarian-zooxanthellar symbiosis

The hydroid Myrionema ambionense, a fast-growing cnidarian (doubling time = 8 days) found in shallow water on tropical back-reefs, lives in symbiosis with symbiotic dinoflagellates of the genus Symbiodinium (hereafter also referred to as zooxanthellae). The symbionts live in vacuoles near the base of host digestive cells, whereas unhealthy looking zooxanthellae are generally located closer to the apical end of the host cell. Cytokinesis of zooxanthellae occurred at night, with a peak in number of symbionts with division furrows (mitotic index, MI = 12%-20%) observed at dawn. The MI of zooxanthellae decreased to near zero by the middle of the afternoon and remained there until the middle of the next night. Densities of live zooxanthellae living inside of host digestive cells peaked following cytokinesis, whereas densities of unhealthy looking symbionts were highest just before the division peak. Mitosis of host digestive cells was highest in the evening, also preceding the peak in zooxanthellar MI. This is the first study relating phased host cell division to diel zooxanthellar division in marine cnidarians. Food vacuoles were prevalent inside of digestive cells of field-collected hydroids within a few hours after sunset and throughout the night, coinciding with digestion of captured demersal plankton. Laboratory experiments showed that food vacuoles appeared in digestive cell cytoplasm within 2 h of feeding with nauplii of Artemia. The number and size of food vacuoles per digestive cell and the percentage of digestive cells with food vacuoles all decreased 5-7 h following feeding in laboratory experiments, and by mid-day in field-collected hydroids. Light and external food supply were important in maintaining phased division of the symbionts, with a lag in response time to both parameters of 11-36 h. Altering light and feeding during the night did not influence the level of the peak MI the next morning, though in one experiment the absence of light slowed final separation of daughter cells at the end of cytokinesis. In another experiment, hydroids starved for 3-7 d and "pulse-fed" Artemia nauplii for 1 h at the beginning of the dark period showed continued low symbiont division (< 5%) after 11 h, whether maintained in constant light or darkness, implying that most algal division is set more than 24 h prior to actual cytokinesis. Transferred to a 14:10 h light:dark cycle for another 24 h (36 h after feeding), the same hydroids exhibited a "normal" peak MI (ca. 15%) at dawn, but zooxanthellae from hydroids kept in constant darkness still showed a low MI. These results show that mitosis of symbiotic dinoflagellates requires three factors: external food; a minimum period of time following feeding (11-36 h), presumably for digestion; and a period of light following feeding, presumably to provide carbon skeletons necessary for completing cytokinesis.     

The effects of tidal currents on the rhythm of feeding and digestion in Pecten maximus L.

The orientation of individuals in two populations of Pecten maximus L., from the west coast of Ireland, shows that they have a marked preference to face directly into a tidal flow. In both localities examined there was a reversal of tides and the members of the populations were divided equally for flood and ebb tides. Twenty-four hour in situ studies of the animals were made and at all stages of the tide individuals facing either due east or west were observed. A cyclical feeding pattern imposed by the reversal of tidal flow is proposed.The pH of various parts of the digestive tract was investigated and showed a wide range of values. The most acid region was that of the stomach. The variations in pH of all of the regions of the gut examined throughout a 24-h period closely followed the pattern of the tidal cycle.Histological analysis of the stomach and digestive diverticula of representative samples of scallops taken at regular intervals over the 24-h periods clearly indicated a diphasic pattern of digestion within the tubules of the digestive diverticula. A close correlation between the phases of intracellular digestion, the pH variations in different regions of the gut, and the tidal cycle indicate distinct feeding cycles in Pecten. Those scallops facing into the ebb current show the same diphasic patterns as those individuals facing the flood current, but are 6 h out of phase. The cycle is considered as a duplication of a diphasic feeding pattern. The tubules themselves undergo a digestive process similar to that in Lasaea rubra (Montagu). The digestive cells phagocytose food material, begin intracellular digestion, and increase in size until they obscure the lumina. The dispersal of waste material and residual bodies is accomplished dramatically by a dehiscence of the tubule cells together with a loss of both digestive and crypt cells. New tubules are regenerated from the apices of those tubules breaking down. In any section of the diverticula tubules in two different conditions are found. The cycle of digestion takes 24 h and in order to facilitate feeding at each 12-h tidal cycle the tubules are equally divided into two phases with one 12 h behind the other.     

Cellular details of the midgut of Cryptocellus boneti (Arachnida: Ricinulei)

The midgut of Cryptocellus boneti was studied by light and electron microscopy. The epithelia of the diverticula and of the anterior part of the midgut tube are composed of two cell types: digestive and secretory. In contrast, the epithelia of posterior part of the midgut tube and of the stercoral pocket consist of one type of cells only. In some places, parts of the midgut system are connected by an intermediate tissue. Digestive cells are characterized by an apical system of tubules, nutritional vacuoles, and spherites; characteristic features of secretory cells are secretory granules and a prominent rough endoplasmic reticulum. Cells of the midgut tube appear not to be involved in the absorption of food. © 1994 Wiley-Liss, Inc.     

TRANSPORT OF UNCONTAMINATED AND MOLLUSCICIDE-CONTAINING FOOD IN THE DIGESTIVE TRACT OF THE SLUG DEROCERAS RETICULATUM (MULLER)

By X-ray analysis of food transport in the alimentary tractof Deroceras reticulatum it was shown that even ten hours afteringestion of a thorium sulfate-containing bait, this materialcan be detected in the crop. After 2.5 h, some parts of labelledfood passes down from the anterior to the posterior part ofthe gut. After 13 h, thorium sulfate-containing material canbe observed only in the gut. After 19 h, no more labelled materialis present in the alimentary tract of the animals. After addition of Cloethocarb, the animals feed on only smallamounts of food. The labelled material only enters the anteriorpart of the gut. After ten hours, the food does not move anymore and does not leave the crop even 19 h after feeding. After molluscicide application, the crop epithelium is moreinfolded than in control animals and the cells are elongated.After 30 h, cells protrude into the lumen of the digestive tract. (Received 11 May 1992; accepted 26 June 1992)     

Structural analysis of the digestive gland of the queen conch Strombus gigas Linnaeus, 1758 and its intracellular parasites

This study describes the structure of the digestive gland ofStrombus gigas in individuals from Guadeloupe and discussesthe function of its cell types and their relationship with intracellularApicomplexa-like parasites. Three cellular types were foundin the epithelium of the blind-ending tubules of the digestivegland according to histological and transmission electron microscopy(TEM) observations; these were: digestive cells, pyramidal cryptcells and vacuolated cells. Columnar digestive cells were characterizedby large Alcian blue-positive granules, which have not beenpreviously described in digestive cells of other caenogastropods.Such granules contain large quantities of proteoglycans thatare exported to the stomach through the physiological destructionof the digestive cells, which undergo a holocrine secretion.Their cytoplasm appears vacuolar due to lipid extraction bysolvents used for tissue preparation. Vacuolated cells alsoappear to be lipid-storage cells. Small triangular-shaped cryptcells, on the other hand, appear to be metabolically activeas suggested by a strong positive in situ hybridization of eukaryoticribosomes, which was confirmed by their large content of ribosomesand rough endoplasmic reticulum compared to the other cell types.These observations suggest that crypt cells may be immaturecells that are involved in the replacement of eliminated digestivecells. However, their spherocrystal inclusions indicate thatthey may be excretory cells or calcium cells. Large brown inclusionswere frequently observed in vacuolated cells; these were identifiedas parasitic protozoans and were present in the digestive glandof all sampled specimens. These protozoans have previously beendescribed from a queen conch population in the San Andres Archipelago(Colombia). Several life cycle stages of the parasite were identifiedby scanning electron microscopy and TEM; trophozoites were characterizedby their conoid-like structure, sporocysts by their thick walls,and gamonts by their thin walls. These observations suggestthat this parasite completes its entire life cycle within thesame host and type of tissue. Although previous investigationsplace this parasite within the Apicomplexa group, further investigationsare necessary in order to confirm the identification of theparasite. (Received 13 May 2008; accepted 3 October 2008)     

THE FINE STRUCTURE OF ACANTHAMOEBA CASTELLANII : I. The Trophozoite

The fine structure of the trophozoite of Acanthamoeba castellanii (Neff strain) has been studied. Locomotor pseudopods, spikelike "acanthopodia," and microprojections from the cell surface are all formed by hyaline cytoplasm, which excludes formed elements of the cell and contains a fine fibrillar material. Golgi complex, smooth and rough forms of endoplasmic reticulum, digestive vacuoles, mitochondria, and the water-expulsion vesicle (contractile vacuole) are described. A canicular system opening into the water-expulsion vesicle contains tubules about 600 A in diameter that are lined with a filamentous material. The tubules are continuous with unlined vesicles or ampullae of larger diameter. Centrioles were not observed, but cytoplasmic microtubules radiate from a dense material similar to centriolar satellites and are frequently centered in the Golgi complex. Cytoplasmic reserve materials include both lipid and glycogen, each of which amounts to about 10% of the dry weight.     

MICROTUBULE ARMS AND CYTOPLASMIC STREAMING AND MICROTUBULE BENDING AND STRETCHING OF INTERTUBULE LINKS IN THE FEEDING TENTACLE OF THE SUCTORIAN CILIATE TOKOPHRYA

Microtubules attached to the pellicle at the tips of tentacles pivot through about 140° on these attachments, splay apart, and bend along their longitudinal axes when feeding occurs. The tubules could be bending in response to pellicular contractions; active bending, sliding, or contraction of the tubules may not be involved. Intertubule links apparently prevent tubules from splaying apart at certain levels. These links are probably under tension during feeding. They stretch; they sometimes become half as thick and eight times as long as they are before feeding. Often, tubules joined together by these links also change in shape; they become slightly flattened and elliptical in cross section. Cytoplasm from the ciliate Tetrahymena is drawn down a feeding tentacle inside an invagination of the Tokophrya cell membrane from the tentacle tip. The positions of arm-bearing microtubules around such invaginations indicate that arms are involved in moving invaginations along. The edges of the perforated Tetrahymena cell membrane are "sealed" to the cell membrane of Tokophrya around each feeding tentacle tip.     

HISTOCHEMISTRY AND ULTRASTRUCTURE OF THE CRYPT CELLS IN THE DIGESTIVE GLAND OF APLYSIA PUNCTATA (CUVIER, 1803)

The crypt cells lining the Aplysia punctata digestive tubulescomprise of three types of cell; calcium, excretory, and thincells. The calcium cells play a role in osmoregulation, mineral storage,exocrine secretion, iron detoxification, and excretion processes.They possess well- developed microvilli and a basal labyrinth,suggesting a role in absorption. The Golgi apparatus is involvedin the production of two main components of calcium spherules;the fibrillar material and mineralized granules. Golgi complex,rough endoplasmic reticulum (RER), ribosomes, and altered mitochondriaare involved in the formation of calcium spherules. Secretoryactivity is indicated by the formation of dense granules containingiron and calcium salts. Lipofuscin pigment has been found inlarge concretions which may arise from cytoplasmic areas surrounded byendoplasmic reticulum, RER and Golgi tubules. There are threestages of excretory cells, called early, mature, and post-excretorycells. This study traces the development of granulofibrillarvacuoles up to the formation of the lipofuscin concretions andshows that excretory cells are in fact degenerating calciumcells. The fine structure of thin cells suggests that they areyoung calcium cells. (Received 29 December 1997; accepted 15 November 1998)     

Ultrastructural studies on the midgut epithelium and digestion in the female tickArgas (Persicargas) arboreus (Ixodoidea: Argasidae)

The ultrastructure of the midgut epithelium and digestion in the female tickArgas (Persicargas) arboreus are described before and after feeding, up to oviposition. The epithelium consists of secretory cells, digestive cells (DI and DII), and regenerative cells which may differentiate into any of the other cell types. In unfed ticks, the midgut wall consists mainly of type DII digestive cells retained from a previous feeding, and a few regenerative cells. Within 3 days after the tick feeding, haemolysis of the host blood components occurs in the midgut lumen. Secretory cells, the first differentiation of the regenerative cells, are presumed to produce a haemolysin and an anticoagulant which are released by merocrine and holocrine secretions. The DII cells seen in unfed ticks, and secretory cells which have completed their secretory cycle, start to have a specialized surface for endocytosis characteristic of type DI digestive cells. From 5 to 7 days after feeding up to the female oviposition, type DI cells which have completed their endocytosis are transformed into type DII digestive cells specialized for intracellular digestion and the storage of reserve nutrients required by the tick for long starvation. The various phases of the digestive cycle are considered according to ultrastructural changes of the midgut epithelium.     

Digestive system membranes: freeze-fracture evidence for differentiation and flow in Paramecium

Freeze-fractured membranes of digestive vacuoles of randomly feeding Paramecium caudatum exhibit dramatic differences in intramembrane particle (IMP) number and distribution on both E- and P-fracture faces. By pulse-feeding latex spheres to cells we have demonstrated that these differences are related to the age of the digestive vacuoles, and that the membranes of such vacuoles undergo a specific sequence of changes during the digestive cycle. Young digestive vacuoles (DV-I; less than or equal to 6 min), nascent vacuoles still connected to the cytopharynx, and discoidal vesicles, from which vacuole membrane is derived, all have a highly particulate E face and a less particulate P face. As early as 3 min after feeding, a second category of digestive vacuoles (DV-II) can be recognized, which are both considerably smaller in diameter and lack particles on their E face. These findings suggest that the endocytic removal of DV-I membrane material associated with the formation of DV-II vacuoles involves a concomitant and selective removal of E-face particles, as essentially no changes are seen in the density of P-face particles on the two types of vacuoles. Beginning at 10 min the first DV-III vacuoles are encountered. These are both larger than the DV-II vacuoles and possess very prominent E-face particles, which resemble those on the E face of the numerous lysosomes bordering the digestive vacuoles. DV-III vacuoles also exhibit a substantial increase in P-face particles. These membrane changes closely parallel, and are probably correlated with, the physiological events occurring within the vacuole lumen: concentration of food, killing of prey, and digestion. Calculations of the amount of membrane removed from DV-I to form DV-II and of the increase in membrane surface area during the transition from DV-II to DV-III indicate that as much as 90% of the initial phagosome (DV-I) membrane can be removed before digestion begins. The enlargment of DV-II must be caused by fusion with adjacent lysosomes which also contribute the new populations of IMPs to the DV- III membrane. The appearance of numerous endocytic structures on older DV-III vacuoles suggests that membrane is retrieved from DV-III before defecation.     

An Ultrastructural Study of Ingestion and Digestion in Tetrahymena pyriformis*

SYNOPSIS. When the structures involved in digestive events in T. pyriformis are examined at the electron microscope level, some information is added to that long known from light microscopy. The food trapping mechanism consists of the three membranelles, undulating membrane, oral ribs, and a “valve” apparently closing the opening to the cytopharynx. Both of the latter structures are supported by microtubules. Fibers extend internally from the cytopharynx and are closely associated with the food vacuole as it forms. Clear vacuoles resembling pinocytic vacuoles appear to arise from differentiated areas of the pellicle and plasma membrane. These vacuoles may fuse with primary lysosomes. Hydrolases are thus contributed to the pinocytic vacuoles which may then fuse with food vacuoles. When first formed food vacuoles contain no hydrolases but may acquire them directly, from primary lysosomes or from pinocytic vacuoles. Digestion proceeds to completion in the food vacuole, at which time soluble food products are released to the cytoplasm. Undigested materials are lost through the cytopyge. In stationary growth phase cells autophagic vacuoles form containing mitochondria and other cellular particulates. Such vacuoles probably contain hydrolases when formed and they may receive others by fusion with primary lysosomes.     

The Ultrastructure of the Digestive Glands in Pinguicula vulgaris L. (Lentibulariaceae) Relative to their Function. I. The Changes During Maturation

The digestive glands of Pinguicula vulgaris become fully maturewhilst still enclosed in the bud. All the gland cells remainintact on the fully expanded unstimulated leaves. As the secretoryhead cells mature, a special layer forms between the plasmalemmaand the cell wall. This layer is shown to be different fromthe typical labyrinthine wall of transfer cells and serves forthe storage of digestive enzymes. Ultrastructural analysis,including morphometry, indicates that the digestive enzymesare synthesized on the RER of the head cells and transferredinto the cell wall, particularly into the slime layer, and vacuoles.This transfer is achieved firstly through continuity of theendoplasmic reticulum with vacuoles (static) and the periplasmicspace (dynamic) and, secondly, into the latter through exocytosisof coated Golgi vesicles and of some vacuoles filled with enzymes. Pinguicula vulgaris L., carnivorous plant, digestive glands, ultrastructure, protein synthesis secretion     

Ultrastructural changes in the gastrodermal phagocytic cells of the planarian Dugesia gonocephala s.l. during food digestion (Plathelminthes)

Summary In the present report the functional morphology of the planarian gastrodermal phagocytic cells is examined in feeding animals. A functional interpretation of some of the morphological findings is given. The events in the fine-structure modifications of the phagocytic cells in the course of phagocytosis and intracellular digestion of food particles were followed through five post-feeding stages in the planarian Dugesia gonocephala. Light and electron microscopical observations demonstrate that there is preliminary intraluminal digestion of food particles; their phagocytosis takes place quickly.Beef hepatocytes that served as food are found engulfed at first in food vacuoles near the apical border of the phagocytic cells, and are clearly recognizable. The vacuoles increase in number to occupy most of the cytoplasm of these cells. Progressive breakdown and disappearance of phagocytosed hepatocytes occurs. In time the vacuoles move deeper into the cells, their contents lose their identity, and condense to homogeneous or heterogeneous residual bodies. These are returned to the distal surface of the cells, and then voided into the intestinal lumen. At the same time, synthesis and accumulation of numerous lipid droplets occurs, probably as a final product resulting from metabolism of the digested material. When feeding is over, the phagocytic cells are filled with lipid droplets, acquiring their typical appearance.It is suggested that disintegration of phagocytic cells during starvation is balanced by proliferation and differentiation of neoblasts into new phagocytic cells during the feeding-starvation cycle.     

On Food Vacuoles in Tetrahymena pyriformis GL

SYNOPSIS. The following problems concerning food vacuoles were studied by in vivo observations of Tetrahymena: (A) Formation of food vacuoles . The process may be divided into 4 stages. Stage 1—gradual growth of the limiting membrane of the open food vacuole (of short duration). Stage 2—"filling up" of the fully expanded vacuole (of long duration). Stage 3—"closing off" of the vacuole (of brief duration). Stage 4—initial movement of the detached vacuole away from the cy-tostome. The possible role of the oral components (apart from membranellar beating) in the process is discussed. (B) Change of pH in the food vacuole . After ingestion of heat-killed yeast stained with indicator dyes (neutral red, bromcresol purple, bromcresol green, bromphenol blue), the observed color changes indicate that pH is neutral in the forming vacuole as well as in newly formed vacuoles; that a pH value of 6.0–5.5 is reached after ∼ 5 min; and that the lowest pH value between 4.0 and 3.5 is reached after 1 hr. Before egestion the pH again increases. (C) Length of the digestive cycle . A determination of the time required to deplete the cells of labeled vacuoles formed during a short exposure, was attempted. Defecation was observed after 1/2 hr and it was frequent after 2 hr. About 25% and 50% of the labeled vacuoles were removed after 1 hr and 2 hr, respectively; however, labeled vacuoles may still be seen in some cells 6 hr after ingestion. The conclusion is that the digestive cycle lasts ∼ 2 hr and that egestion of undigestible material is a random process.     

EVOLUTION, DURING DIGESTION, OF THE BACTERIAL FLORA IN THE ALIMENTARY SYSTEM OF HELIX ASPERSA (GASTROPODA: PULMONATA): A SCANNING ELECTRON MICROSCOPE STUDY

The use of scanning electron microscopy (SEM) allowed a studyof the distribution of bacteria in the various digestive organsof the snail Helix aspersa Müller. The bacteria are enclosedby mucous secretions (mucous film or mucous grains) and sometimesattached on the cilia of some of the digestive walls. Accordingto the food that was given to the snails, different morphologicaltypes appeared, two of which dominated. Adult snails were fasted for 4 days, given a dehydrated artificialfood and then sacrificed at different times during digestion.The presence of bacteria may be related to the time of digestion.In fact, bacteria seem to accompany the food mass; they developmostly in the stomach and in the intestine where they may helpdigest the food. Fasting or hibernating snails do not possess bacteria in thealimentary lumen or on the digestive walls. However, the residualfaeces localized in the distal, intestinal lumen, lodge greatquantities of bacteria. From these results, the endogenous or/and exogenous existenceof the bacterial flora in alimentary system of Helix aspersais discussed. (Received 26 June 1989; accepted 16 October 1989)     

Some Aspects of Nutrition in Parasites

Some selected examples of food-getting mechanisms in animalparasites are reviewed. The range of adaptations includes: (1)mechanical devices for biting or sucking plus internal or externaldigestive capacity, (2) internal digestive capacities only,or (3) no mechanical feeding mechanisms and no digestive capacities.In the latter case, feeding mechanisms are restricted to absorptivecapacity. Available data on absorption are briefly reviewed. Questions concerning the food of parasites in hosts are posed,and the following examples are analyzed to demonstrate the difficultiesin answering these questions: Fasciola, hookworms, Trichuris,and some nematode parasites of vertebrates.     

Developmental Features of the Primary Phloem in Phaseolus vulgaris L.

Certain developmental features of the primary phloem were examinedin Phaseolus vulgaris L., chiefly by the use of the pulvinusat the base of the petiole. The cells included in the studywere the sieve element, the companion cell, and the tannin cell.In the sieve element, the sieve plate shows the usual sequenceof conversion of plasmodesmatal canals into pores. The endoplasmicreticulum, which appears as flat cisternae associated with ribosomesin younger cells, later becomes in part stacked and in partaligned parallel with the walls as a network. The stacked ERprecedes the anastornosing parietal ER in time of development,but the parietal ER persists longer. Of the two forms of P-proteincharacteristic of a number of Fabaceae, the crystalline bodyappears considerably in advance of the body composed of tubules.Neither form of P-protein disperses completely in the maturecell, although the crystalline protein may spread out into aggregatesof fine fibrils. The companion cells show the typical denseprotoplasts and branched plasrnodesmatal connections with thesieve elements. The vacuome of these cells is dispersed intonumerous small vacuoles, many of which appear to be concernedwith autophagic digestion of protoplasmic material. The tannincells have large vacuoles in which the tannin material is located.The cells form vertical series in which the end walls becomeperforated.