THE BIOLOGY OF HARVEYELLA MIRABILIS (CRYPTONEMIALES,RHODOPHYCEAE). VI. TRANSLOCATION OF PHOTOASSIMILATED 14C122

Pulse-chase labelling experiments demonstrate that photoassimilated ¹⁴C-bicarbonate is translocated from the host red alga Odonthalia floccosa (Esper) Falkenberg to the parasite Harveyella mirabilis (Reinsch) Schmitz & Reinke. The primary path of translocation is from host cortical cells (the site of photoassimilation) to the erumpent parasite pustule via the zone of interdigitation. The latter is a tissue region in which rhizoidal cells of Harveyella grow between, and establish secondary pit plugs with medullary cells of Odonthalia. A secondary translocation pathway occurs from isolated host cells dispersed in the pustule of Harveyella to adjacent parasite cells.     

THE EVOLUTION OF PARASITISM IN RED ALGAE: CELLULAR INTERACTIONS OF ADELPHOPARASITES AND THEIR HOSTS1

In the initial stages of cell–cell interactions (spore germination and host penetration), the adelphoparasites Gardneriella tuberifera Kyl. and Gracilariophila oryzoides Setch. & Wilson form infection rhizoids that fuse directly with underlying host epidermal or cortical cells. In so doing, parasite nuclei and other organelles enter the cytoplasm of the host. The resulting heterokaryon may fuse with adjacent host cells either directly, via secondary pit connections, or by the dissolution or dislodgment of pit plugs from existing pit connections. The cell fusion events result in a heterokaryotic syncytium in which parasite nuclei replicate. In Gardneriella, formation of the syncytium induces surrounding host tissues to divide to form a photosynthetic callus. The internalized syncytium forms conjunctor and rhizoidal cells that fuse with host callus, eventually transforming the host callus into cells containing parasite nuclei. Gracilariophila does not induce surrounding host tissue to divide. Rather, division of the initial heterokaryotic tissue gives rise to the colorless mantle that protrudes from the host and forms reproductive structures. The heterokaryotic tissue also fuses with underlying host cells, thereby spreading parasite nuclei throughout adjacent host cells. In both these adelphoparasites, transformation of host cells by parasite nuclear invasion results in plastid dedifferentiation, an increase in mitochondria, autolysis of organelles, and accumulation of large amounts of floridean starch. The development and physiology of these parasites is similar to normal post-fertilization processes in the hosts that give rise to carposporophytes and suggests that these adelphoparasites may have originated from perturbations of developmental pathways involved in their host's post-fertilization development.     

The structure and formation of host-parasite pit connections between the red algal alloparasiteHarveyella mirabilis and its red algal hostOdonthalia floccosa

Summary Harveyella mirabilis is a colourless red algal alloparasite which grows on and within its photosynthetic hostOdonthalia floccosa. Cells ofHarveyella establish secondary pit connections (PCs) with other parasite cells and with cells of the host. Small, uninucleate conjunctor cells are produced by parasite cells and remain connected to them by PCs. Conjunctor cells may fuse with either an adjacent host or parasite cell, with the parasite-conjunctor cell PC becoming either a host-parasite or parasite-parasite secondary PC. Occasionally the conjunctor cell does not fuse with an adjacent cell (either host or parasite) and degenerates. The secondary pit plug which forms between a parasite cell and its conjunctor cell always develops with two structurally distinct surfaces characteristic of a host-parasite pit plug. Only if the conjunctor cell fuses with another parasite cell will the structure of the pit plug be altered to that of a parasite-parasite pit plug. Fungal hyphae also invade the region of infection, andHarveyella cells respond by producing nonfunctional conjunctor cells that grow towards adjacent hyphae. Evidence suggests that secondary PCs may be induced to form mechanically, by the physical presence of another cell, rather than in direct response to a message received from an adjacent cell. The mechanism of secondary PC formation described here is similar to that reported for the closely related alloparasiteHolmsella and may be common to a number of red algal parasitic associations. Helen Margaret Quirk, B. Sc. (Hons), M. Sc. (1953–1982), student, research assistant and friend, died after a long illness on October 24, 1982.     

LEACHIELLA PACIFICA,GEN. ET SP. NOV., A NEW PARASITIC RED ALGA FROM WASHINGTON AND CALIFORNIA

Leachiella pacifica, gen. et sp. nov., a marine alloparasitic red alga is described from Washington and California. Several species of Polysiphonia and Pterosiphonia are hosts for this parasite. The thallus is a white, multiaxial, unbranched pustule with rhizoidal filaments that ramify between host cells, forming numerous secondary pit connections with host cells. All reproductive structures develop from outer cortical cells. Tetrasporocytes, situated on stalk cells, undergo simultaneous, tetrahedral cleavage to form tetraspores. Spermatia are formed continuously by oblique cleavages of the elongate spermatial generating cells. This results in spermatial clusters consisting of 4–8 spermatia in an alternate arrangement. Carposporophyte development is procarpial. The carpogonium is part of a six-celled branch including a sterile cell that is formed by the basal cell. The carpogonial branch is attached laterally to an obovate supporting cell that also forms an auxiliary cell, presumably formed prior to fertilization. After fertilization the carpogonium temporarily fuses with the auxiliary cell apparently to transfer the diploid nucleus and initiate further fusion with the subtending supporting cell to form an incipient fusion cell. The auxiliary cell portion of this fusion cell divides to form gonimoblast initials that continue to divide, forming gonimoblast filaments whose terminal cells differentiate into carpospores. The remainder of the fusion cell enlarges by continual fusion with adjacent vegetative cells. The resultant carposporophyte consists of a basal, multinucleate fusion cell supporting a hemispherical cluster of gonimoblast filaments with terminally borne carpospores. Vegetatively, Leachiella resembles several other parasitic red algae but it is clearly separated by the procarp, carposporophyte development and structure, and tetrasporocyte cleavage.     

THE ROLE OF SECONDARY PIT CONNECTIONS IN RED ALGAL PARASITISM1

Secondary pit connections are common between cells of hosts and parasites in the widespread phenomenon of red algal parasitism. The DNA-specific fluorochrome 4′,-6-diamidino-2-phenylindole (DAPI) reveals that in host-parasite secondary pit connection (SPC) formation between the parasitic red alga Choreocolax polysiphoniae and its host Polysiphonia confusa, a nucleus and other cytoplasmic components of the parasite are delivered into the cytoplasm of a host cell. Host cells receive large numbers of parasite nuclei and these, apparently arrested in G¹, are maintained intact in host cells for periods of several weeks. Within these enlarged, differentiated cells, starch accumulates and cytoplasmic organelles proliferate as the central vacuole decreases in size. Host nuclear DNA synthesis is stimulated in the infected host cell, resulting in an increase in the number of host nuclei, or an increase in DNA in each of the existing host nuclei (i.e. somatic polyploidy). Occasionally, infected host cells will recommence division and engender a new host branch. Microspectrofluorometry of nuclear DNA quantitatively confirms not only the identity and transfer of parasite nuclei to host cells, but also the transfer of parasite nuclei to other parasite cells. Measurements also reveal that the single nucleus of Choreocolax becomes progressively more polyploid as cells become larger and more highly differentiated. Secondary pit connection formation between Choreocolax and Polysiphonia provides the mechanism for the transfer of parasite genetic information (via the parasite nucleus and cytoplasm) into the host. The parasite nuclei may thereby control and redirect the physiology of the host for the benefit of the parasite.     

The fine structure and cytology of the association between the parasitic red algaHolmsella australis and its red algal hostGracilaria furcellata

Summary Holmsella australis Noble andKraft ms. is a colourless red algal parasite, forming whitish pustules on its photosynthetic red algal host,Gracilaria furcellata Harvey. In the infected region, host cortical tissue continues to grow and enclose the expanding pustule. Filaments of both host and parasite grow apically, the cells being connected by primary pit connections (PCs). Secondary PCs form between cells of the same species, and in addition,H. australis initiates the formation of secondary PCs with cells ofG. furcellata. All three types of secondary PC are morphologically distinct. In hostparasite PCs the surface adjoining the host cell is similar in structure to a host-host PC, while that adjoining the parasite cell has the structure of a parasite-parasite PC. The plasma membrane is continuous between the cells of the unrelated host and parasite. In addition, a cap membrane is typically produced only on the host surface, though occasionally the parasite side is enclosed by a cap membrane as well. Cap membranes are absent from parasite-parasite PCs (making them intracellular), while host-host PCs are typically extracellular, both cells producing cap membranes. The presence or absence of a cap membrane in certain positions appears to vary, and suggests that cells may be able to regulate its presence. Since transport of nutrients would be expected to occur from host to parasite cells, and between parasite cells, the morphological evidence presented here suggests the PCs may be the pathway.     

The fine structure of secondary pit connection formation between the red algal alloparasiteHolmsella australis and its red algal hostGracilaria furcellata

Summary Pit connections (PCs) develop between the parasitic red algaHolmsella and its hostGracilaria. Only parasite cells initiate the formation of host-parasite pit connections. The parasite produces a small connecting cell (termed the conjunctor cell) which moves through the cell wall to fuse with either an adjacent host or parasite cell. The parasite secondary PC, which forms between the conjunctor cell and the parasite cell, is structurally different from a parasite primary PC, and has the distinct structure of a host-parasite PC. Only if the conjunctor cell fuses with another parasite cell will the former parasite-conjunctor cell PC be altered to a typical parasite-parasite PC. If the conjunctor cell fuses with an adjacent host cell the PC continues to develop as host-parasite. Occasionally a conjunctor cell fails to fuse with an adjacent cell (whether host or parasite), and the conjunctor cell and PC eventually breakdown in the cell wall. The parasite overcomes several barriers in order to infect the host, including the formation of host-parasite PCs which appear to be a necessary component of the parasiticHolmsella-Gracilaria association.     

THE BIOLOGY OF HARVEYELLA MIRABILIS (CRYPTONEMIALES,RHODOPHYCEAE). VII. STRUCTURE AND PROPOSED FUNCTION OF HOST-PENETRATING CELLS1

The parasitic red alga Harveyella mirabilis (Reinsch) Schmitz & Reinke was examined by light and electron microscopy to determine the structural mechanism involved in nutrient transfer. The host-penetrating rhizoidal cells are unique in possessing an extensive and apparently dynamic endomembrane system as well as other unique cytoplasmic inclusions. The membrane system consists of the plasmalemma, pinocytotic vesicles, multivesicular and concentric bodies, endoplasmic reticulum, dictyosomes, micro-body-like structures and an extensive vacuolar system. It is proposed that this system is active in the uptake and processing of host-derived nutrients. Plasmalemmal extensions (plasmalemmavilli) of Harveyella medullary cells may also function in nutrient uptake.     

Development of the red algal parasite Vertebrata aterrimophila sp. nov. (Rhodomelaceae,Ceramiales) from New Zealand

Parasitic red algae grow only on other red algae and have over 120 described species. Developmental studies in red algal parasites are few, although they have shown that secondary pit connections formed between parasite and host and proposed that this was an important process in successful parasitism. Furthermore, it was recorded that the transfer of parasite nuclei by these secondary pit connections led to different host cell effects. We used developmental studies to reconstruct early stages and any host cell effects of a parasite on Vertebrata aterrima. A mitochondrial marker (cox1) and morphological observations (light and fluorescence microscopy) were used to describe this new red algal parasite as Vertebrata aterrimophila sp. nov. Early developmental stages show that a parasite spore connects via secondary pit connections with a pericentral host cell after cuticle penetration. Developmental observations revealed a unique connection cell that grows into a ‘trunk-like’ structure. Host cell transformation after infection by the parasite included apparent increases in both carbohydrate concentrations and nuclear size, as well as structural changes. Analyses of molecular phylogenies and reproductive structures indicated that the closest relative of V. aterrimophila is its host, V. aterrima. Our study shows a novel developmental parasite stage (‘trunk-like’ cell) and highlights the need for further developmental studies to investigate the range of developmental patterns and host effects in parasitic red algae.     

PARASITIC INTERACTIONS AND VEGETATIVE ULTRASTRUCTURE OF CHOREOAEMA THURETII (CORALLINALES,RHODOPHYTA)1

The monotypic coralline red alga, Choreonema thuretii (Bornet) Schmitz (Choreonematoideae), grows endophytically within three geniculate genera of the Corallinoideae. Although the thallus of Choreonema is reduced, lacks differentiated plastids, and is endophytic except for its conceptacles, its status as a parasite has been questioned because cellular connections to the host had not been ob served. Transmission electron microscopy, however, disclosed a previously undescribed type of parasitic interaction in which Choreonema interacts with its host through specialized cells known as lenticular cells. These small, lens-shaped cells are produced from the single file of host-penetrating vegetative cells. Pit plug morphology between vegetative and lenticular cells is polarized. Plug caps facing the vegetative cell have normal coralline morphology, while those facing the lenticular cell are composed of three layers. Regions of lenticular cells near host cells protrude toward the host cell; upon encountering the host cell wall, the prolrusion produces numerous finger-like fimbriate processes that make cellular connections with the host cell. Lenticular cells may extend several protrusions toward a host cell or penetrate more than one host cell; two or more lenticular cells may also penetrate the same host cell. The lack of secondary pit connections, cell fusions, and passage of parasitic nuclei suggest that this parasitic relationship may be evolutionarily older than previously reported cases of parasitism in red algae.     

COMPARATIVE DEVELOPMENT OF THE RED ALGAL PARASITES BOSTRYCHIOCOLAX AUSTRALIS GEN. ET SP. NOV. AND DAWSONIOCOLAX BOSTRYCHIAE (CHOREOCOLACACEAE,RHODOPHYTA)1

The development of two red algal parasites was examined in laboratory culture. The red algal parasite Bostrychiocolax australis gen. et sp. nov., from Australia, originally misidentified as Dawsoniocolax bostrychiae (Joly et Yamaguishi-Tomita) Joly et Yamaguishi-Tomita, completes its life history in 6 weeks on its host Bostrychia radicans (Montagne) Montagne. Initially the spores divide to form a small lenticular cell, and then a germ tube grows from the opposite pole. Upon contact with the host cuticle, the germ tube penetrates the host cell wall. The tip of the germ tube expands, and the spore cytoplasm moves into this expanded tip. The expanded germ tube tip becomes the first endophytic cell from which a parasite cell is cut off that fuses with a host tier cell. The nuclei of this infected host cell enlarge. As parasite development continues, other host-parasite cell fusions are formed, transferring more parasite nuclei into host cells. The erumpent colorless multicellular parasite develops externally on the host, and reproductive structures are visible within 2 weeks. Tetrasporangia are superficial and cruciately or tetra-hedrally divided. Spermatia are formed in clusters. The carpogonial branches are four-celled, and the carpogonium fuses directly with the auxiliary (support) cell. The mature carposporophyte has a large central fusion cell and sympodially branched gonimoblast filaments. Early stages of development differ markedly in Dawsoniocolax bostrychiae from Brazil. Upon contact with the host, the spore undergoes a nearly equal division, and a germ tube elongates from the more basal of the two spore cells, penetrates the host cell wall, and fuses with a host tier cell. Subsequent development involves enlargement of the original spore body and division to form a multicellular cushion, from which descending rhizoidal filaments form that fuse with underlying host cells. This radically different development is in marked contrast to the final reproductive morphology, which is similar to B. australis and has lead to taxonomic confusion between these two entities. The different spore germination patterns and early germ-ling development of B. australis and D. bostrychiae warrant the formation of a new genus for the Australian parasite.     

LAURENCIA MARILZAE SP. NOV. (CERAMIALES,RHODOPHYTA) FROM THE CANARY ISLANDS,SPAIN, BASED ON MORPHOLOGICAL AND MOLECULAR EVIDENCE1

Laurencia marilzae Gil‐Rodríguez, Sentíes et M.T. Fujii sp. nov. is described based on specimens that have been collected from the Canary Islands. This new species is characterized by distinctive yellow–orange as its natural habitat color, a terete thallus, four pericentral cells per vegetative axial segment, presence of secondary pit‐connections between adjacent cortical cells, markedly projecting cortical cells, and also by the presence of corps en cerise (one per cell) present in all cells of the thallus (cortical, medullary, including pericentral and axial cells, and trichoblasts). It also has a procarp‐bearing segment with five pericentral cells and tetrasporangia that are produced from the third and fourth pericentral cells, which are arranged in a parallel manner in relation to fertile branchlets. The phylogenetic position of this taxon was inferred based on chloroplast‐encoded rbcL gene sequence analyses. Within the Laurencia assemblage, L. marilzae formed a distinctive lineage sister to all other Laurencia species analyzed. Previously, a large number of unique diterpenes dactylomelane derivatives were isolated and identified from this taxon. L. marilzae is morphologically, genetically, and chemically distinct from all other related species of the Laurencia complex described.     

The Endophytic System of Arceuthobium minutissimum, the Indian Dwarf Mistletoe

The Indian dwarf mistletoe, Arceuthobium minutissimum Hook f.is the most diminutive dicotyledonous stem parasite on Pinusexcelsa. The endophytic system is well developed, having a largenumber of anastomosing strands in the cortex and sinkers penetratingthe medullary rays in wood. The cortical strand is protostelicwith the central tracheary elements, the vessels, surroundedby paren-chymatous cells. An earlier report of absence of vesselsseems to be erroneous. The growth of the cortical strands iseffected by an apical cell. The sinkers typically associatedwith the rays of host, are composed of parenchymatous cellsand tracheary elements including vessels. They make contactswith the cells of the ray through pits present in the trachearyelements. The sinkers cause hypertrophy and even fusion of twoor more rays to form a composite medullary ray. The tracheidsof the host tissue also become stunted and contorted in shape.These observations are in agreement with those of other investigatorson American host species for Arceuthobium.     

DEVELOPMENT OF JANCZEWSKIA MORIMOTOI (CERAMIALES) ON ITS HOST LAURENCIA NIPPONICA (CERAMIALES,RHODOPHYCEAE)1

Janczewskia morimotoi Tokida was successfully cultured from spore to reproductive maturity on its host Laurencia nipponica Yamada. The spore penetrates the host without requirement for wound or abrasion sites, growing between host cortical cells and developing a superficial and an endophytic system simultaneously. During the juvenile period, when the parasite is nonpigmented, it differentiates a cortex and the proliferating endophytic filaments enlarge causing a displacement of layers of host cells into the parasitic tissue. Host cells contacted by cells of the parasite exhibit increased wall thickness, cytoplasmic density and vesicle formation. Pit connections between host and parasite cells were rarely observed whereas penetration of host cell walls was seen commonly. As the parasite increases in size, its cells become pigmented evenly throughout the cortex and host cells show less obvious reactions to the parasite. At this same time, the parasite develops branches and reproductive structures. Host plant segments less than 3 cm long failed to grow when infected with spores of the parasite whereas longer segments were not significantly affected by the parasite. In the absence of the host, the parasite cannot complete its development. Although J. morimotoi is well pigmented at maturity, the absence of pigmentation in the juvenile stage, penetration of host cells, and effect on host growth in culture strongly suggest that it is parasitic during at least its early development.     

Drachiella liaoii sp. nov., a new member of the Schizoserideae (Delesseriaceae,Rhodophyta) from Taiwan and the Philippines

A new member of Delesseriaceae (Ceramiales, Rhodophyta) is described from Southern Taiwan and the Philippines. On the basis of comparative vegetative and reproductive morphology, and phylogenetic analysis inferred from nuclear-encoded large-subunit ribosomal DNA sequences (LSU rDNA), we conclude that it belongs in the genus Drachiella, tribe Schizoserideae, subfamily Phycodryoideae. The new taxon shares with other Drachiella species the absence of macro- and microscopic veins; diffuse growth by marginal and intercalary meristematic cells; a polystromatic, lobed thallus; abundance of rhizoidal marginal proliferations used for attachment; convoluted plastids in surface cells; abundant secondary pit connections among adjacent vegetative cells; large intercellular spaces between surface cells; procarps confined to the upper side of the thallus, circular in outline, consisting of a supporting cell bearing a strongly curved carpogonial branch and two sterile groups that remain undivided; vertical division of gonimoblast initial from auxiliary cell, and unilateral, monopodial branching of gonimoblasts; and mature cystocarps with a massive candelabrum-like fusion cell of fused gonimoblasts bearing carposporangia in branched chains. It is distinguished from the other members of the genus by thalli that consist of extensive tangled mats of prostrate and overlapping decumbent blades, procarps confined to the upper side of the thallus, and the lack of basal stalks or stipes. Whereas the Schizoserideae is predominantly a Southern Ocean tribe, one of the tribe's four genera, Drachiella, was known only from the eastern Atlantic and Mediterranean. We herein report the first record of the genus for the Indo-Pacific Ocean, and describe Drachiella liaoii, sp. nov., as a fourth species in the genus.     

CONCEPTACLE STRUCTURE OF THE PARASITIC CORALLINE RED ALGA CHOREONEMA THURETII (CORALLINALES) AND ITS TAXONOMIC IMPLICATIONS1

The major diagnostic features for erecting the red algal subfamily Choreonematoideae (Corallinales) were a combination of 1) absence of both cell fusions and secondary pit connections, 2) conceptacle roof and wall comprised of a single cell layer, and 3) presence of tetrasporangial pore plugs within a uniporate conceptacle in the monotypic taxon Choreonema thuretii (Bornet) Schmitz. Because this alga is a parasite, the absence of secondary cell connections is most likely an adaptation to a reduced thallus. This study shows that all conceptacles are not composed of a file of cells but rather a single layer of epithallial cells that are underlain by a thick layer of calcified acellular material; both epithallial cells and the calcified layer are produced by peripheral sterile cells. Although the outermost tetrasporangial pore canal is uniporate, there is a calcified acellular multiporate plate recessed just below the rim. The plate is produced by interspersed sterile cells and is continuous with the calcified layer supporting the conceptacle. These unique structures are likely due to parasitism rather than to the ancestral state. Based on these results and a reexamination of published micrographs depicting lenticular cells in Austrolithon intumescens Harvey et Woelkerling, we propose that both subfamily Choreonematoideae and Austrolithoideae are closely allied with subfamily Melobesioideae. This distant relationship to its host (Corallinoideae) plus a combination of unique conceptacle and unusual type of parasitism indicates that C. thuretii is an alloparasite and that it is likely the most ancient red algal parasite studied to date.     

Comparative morphology and systematics of Chondrymenia lobata from the Mediterranean Sea and a phylogeny of the Chondrymeniaceae fam. nov. (Rhodophyta) based on rbcL sequence analyses

The Chondrymeniaceae Rodríguez-Prieto, G. Sartoni, S.-M. Lin & Hommersand, fam. nov., is proposed for Chondrymenia lobata. Analyses of rbcL sequences place the new family in a large gigartinalean assemblage that comprises the Cystocloniaceae–Solieriaceae complex. Plants are decumbent and growth takes place by division of multiple apical cells at the margin of the blade. Thalli consist of an outer cortex of subspherical to elongate cortical cells arranged in anticlinal rows, a subcortex of cells cross-linked by lateral arms, and a large central medulla composed of primary medullary filaments intermixed with numerous rhizoidal filaments. Male stages are reported in monoecious individuals. Inactive carpogonial branches consist of a two-celled filament that is directed inwards from the supporting cell. Functional carpogonial branches are oriented outwardly, with the carpogonia and trichogynes pointed towards the thallus surface. After presumed fertilization, the carpogonium fuses with the hypogynous cell and transfers the zygote nucleus. The hypogynous cell, in turn, fuses with the supporting cell which contains many haploid nuclei. The resulting fusion cell functions as an auxiliary cell that cuts off a single gonimoblast initial, which produces the gonimoblast filaments. Gametophytic cells close to the auxiliary cell unite with it to form a placental fusion network of variable size and outline, and a placental fusion cell. Proximal gonimoblast cells fuse with the placental fusion cell, while the distal cells differentiate into branched chains of subspherical carposporangia. The superficial similarity of the outwardly developed osteolate cystocarp is responsible for Kylin's (1956) placement of Chondrymenia in his family Sarcodiaceae; however, the manner in which the placenta is formed is more like that seen in the Cystocloniaceae–Solieriaceae complex.     

Lithophyllum cuneatum sp. nov. (Corallinaceae, Rhodophyta), a new species of non-geniculate coralline alga semi-endophytic in Hydrolithon onkodes and Neogoniolithon sp. from Fiji, South Pacific

A new species of semi-endophytic coralline alga, Lithophyllum cuneatum (Corallinaceae: Lithophylloideae), is described from Fiji. The species is characterized by a wedge-like thallus that is partially buried in the thallus of the host coralline, Hydrolithon onkodes (Heydrich) Penrose et Woelkerling or occasionally Neogoniolithon sp., and that appears at the surface of the host as a small pustule that is usually paler in color than the host. The thallus consists of erect filaments that are derived from a single cell. The basal cell, when visible, is non-palisade, and areas of bistratose margin are absent. Cells of contiguous erect filaments are joined by secondary pit connections. Epithallial cells are present in 2–3 layers, and individual trichocytes are common. Gametangial plants are dioecious. Male conceptacles have simple spermatangial systems that are confined to the floors of their elliptical chambers. Carposporangial conceptacles contain 5–8 celled gonimoblast filaments that are borne at the margin of a more-or-less discoid fusion cell, and so occupy the periphery of the elliptical conceptacle chambers. Tetrasporangial conceptacles are uniporate, with roofs formed from peripheral filaments, and chambers lack a central columella of sterile filaments. Despite its semi-endophytic nature, haustorial cells are absent, and plastids and pigmentation are present.     

Life Cycle of <Emphasis Type="Italic">Plasmodiophora brassicae</Emphasis>

Plasmodiphora brassicae is a soil-borne obligate parasite. The pathogen has three stages in its life cycle: survival in soil, root hair infection, and cortical infection. Resting spores of P. brassicae have a great ability to survive in soil. These resting spores release primary zoospores. When a zoospore reaches the surface of a root hair, it penetrates through the cell wall. This stage is termed the root hair infection stage. Inside root hairs the pathogen forms primary plasmodia. A number of nuclear divisions occur synchronously in the plasmodia, followed by cleavage into zoosporangia. Later, 4–16 secondary zoospores are formed in each zoosporangium and released into the soil. Secondary zoospores penetrate the cortical tissues of the main roots, a process called cortical infection. Inside invaded roots cells, the pathogen develops into secondary plasmodia which are associated with cellular hypertrophy, followed by gall formation in the tissues. The plasmodia finally develop into a new generation of resting spores, followed by their release back into soil as survival structures. In vitro dual cultures of P. brassicae with hairy root culture and suspension cultures have been developed to provide a way to nondestructively observe the growth of this pathogen within host cells. The development of P. brassicae in the hairy roots was similar to that found in intact plants. The observations of the cortical infection stage suggest that swelling of P. brassicae-infected cells and abnormal cell division of P. brassicae-infected and adjacent cells will induce hypertrophy and that movement of plasmodia by cytoplasmic streaming increases the number of P. brassicae-infected cells during cell division.     

TRANSLOCATION OF 11C-PHOTOASSIMILATE IN THE BLADE OF MACROCYSTIS PYRIFERA (PHAEOPHYCEAE)1

Radioactive bicarbonate was pulse fed to blades of Macrocystis pyrifera (L.) C. A. Ag. and the movement of the ¹¹C-labelled photoassimilates was monitored in vivo using an externally mounted array of Geiger-Müller detectors. Results of experiments conducted in August 1982 and February 1983 showed kinetic transport profiles composed of short pulses of ¹¹C (periods of two to three minutes and six to eight minutes) and a mass flow component travelling with a speed of 6–22 cm · h^?1. The pulse-like movement of ¹¹C-photoassimilates, revealed for the first time in a kelp, may be driven by an energy-assisted transport mechanism. Light microscopy revealed a putative symplastic transport pathway from the photo synthetic meristoderm to the medullary sieve cells in the M. pyrifera blade. Of particular importance were the connections between the inner cortical cells and thin-walled medullary sieve cells. Electron microscopy showed sieve plate pore diameters ranging between 35–60 nm in the cortex and ca. 40 nm in the end walls of the thin-walled sieve cells.