Spiral growth in the radially-expanding piloboloid mutants ofPhycomyces blakesleeanus

The growth zone of the sporangiophore of a piloboloid mutant,pil, ofPhycomyces expands radially at an increased rate until the growth zone becomes nearly spherical, in sharp contrast to that of the wild-type sporangiophore which exhibits longitudinal elongation only and is conical. The rotation of thepil sporangiophore reverses its direction from clockwise (CW) to counterclockwise (CCW) during the period of increased radial expansion, and the CCW rotation continues as long as does the radial expansion. The direction of rotation and the time of reversal are correlated with the relative rates of cell-wall expansion in the longitudinal and transverse directions. The CCW rotation of the sporangiophore of this mutant can be explained by the behavior of the microfibrils, as previously proposed to explain the rotation of the wild-type sporangiophore.Abbreviations CW clockwise - CCW counterclockwise — both as viewed from above     

Effects of microbeam light on growth and phototropism ofPilobolus crystallinus sporangiophores

A small blue-light beam (50 μm in diam) was used to examine light-growth response and phototropism inPilobolus crystallinus sporangiophores. Continuous irradiation by microbeam of a region 100–150 μm from the apex promoted the growth of a dark-adapted sporangiophore for about 15 min after a lag period of 1–2 min. After the promotion, the growth rate fell below that before the irradiation. Irradiation of the apex of sporangiophore slightly promoted the growth but strongly inhibited the growth after the promotion. A smaller light beam (10 μm in diam) applied continuously at grazing incidence along one side of the sporangiophore caused bending toward the shaded side, implying that the irradiated side grew more rapidly than the shaded side and that the lens effect is involved in the phototropism of young sporangiophores ofP. crystallinus. The involvement of the lens effect was confirmed by the fact that a carotenoid-less mutant was 1.5–2 times more sensitive to unilateral blue light than the wild type, probably because of a smaller intracellular light attenuation during passage through the mutant cell.     

Ethylene is involved in the autochemotropism of Phycomyces

The sporangiophore of the fungus Phycomyces blakesleeanus has the property of growing away from a barrier which is few mm from the growing zone of the sporangiophore (avoidance or autochemotropic response). A model has been published (Cohen, R.J., Jan, N.Y., Matricon, J., Delbrück, M.: J. Gen. Physiol. 66, 67–95 (1975)). To explain the avoidance response which postulates that the sporangiophore emits and readsorbs a volatile growth-promoting effector (gas X) and that the barrier modifies the effector distribution by acting as an aerodynamic obstacle, causing a higher concentration of gas X on the side of the sporangiophore closer to the barrier. From this model we deduced three properties of the gas X. Of the several gases tested (N₂, CO₂, CH₄, C₂H₂, C₂H₄, C₂H₆) only ethylene (C₂H₄) had all these three properties, a finding which suggests that it has a role in the avoidance response (autochemotropism).Abbreviation Spph Sporangiophore     

Inhibition of phototropism inPhycomyces sporangiophores by calmodulin antagonist and antimicrotubular agents

A simple method was devised to measure growth and phototropism in isolated sporangiophores fromPhycomyces blakesleeanus. With this method, the effects of a number of substances that affect different metabolic reactions were tested. A large number of compounds inhibited or stimulated sporangiophore growth, but did not affect the phototropic response. Only colchicine, thiabendazole, and specially trifluoperazine affected both sporangiophore growth and phototropism. These results suggest that microtubules and calmodulin are involved in the response of the fungus to light.     

Analysis of growth and rotational behavior of sporangiophores in Pilobolus crystallinus (Wiggers) Tode

The growth and rotation of the sporangiophore of Pilobolus crystallinus, which are important factors in its phototropic behavior, were analyzed throughout its development. The sporangiophore initial emerged from the trophocyst and elongated at the extreme tip without rotating. The elongation rate of the sporangiophore apex then gradually decreased and the apex expanded radially to produce the sporangium, but no rotation occurred. A transient cessation of elongation after sporangium development was followed by resumption of both elongation and radial expansion in the region beneath the sporangium developing the subsporangial vesicle. Rotation was not obvious at this stage. Radial expansion of the subsporangial vesicle continued at a decreasing rate until full size was reached. Elongation then recommenced in the newly established growth zone in the upper region of the sporangiophore just beneath the subsporangial vesicle. During this period of growth, the sporangiophore rotated in a clockwise direction as viewed from above. All growth and rotation ceased about 1 h before ejection of the sporangium into the air. Based on these results, a modified classification of the developmental stages has been proposed.This work was carried out under the Joint Research Program of the Institute of Genetic Ecology, Tohoku University, Japan (892006). The authors please to thank Kaori Koga and Hiroko Kikuchi for their helpful assistance.     

Effects of light and temperature on sporangiophore initiation in Pilobolus crystallinus (Wiggers) Tode

Sporangiophore initiation in Pilobolus crystallinus grown in white light was induced by either a dark or a low-temperature treatment. The period of darkness necessary to induce sporangiophore initiation was shortened by lowering the temperature. Arrhenius plots for the sporangiophore-suppressing reaction in both light and darkness consisted of two straight lines with a Q₁₀ of about 2 at lower temperatures and 8–11 at higher temperatures. The temperature at which the Q₁₀ changed was the lower, the higher the fluence rate: 14° C at 8 W/m², 19.5° C at 0.24 W/m² and 24.5° C in darkness. Possible interpretations of these results are briefly presented.Abbreviations %SP percentage of trophocysts initiating sporangiophores - D50% duration of treatment required to 50% sporangiophore initiation     

The sensitive period for light and temperature regulation of sporangiophore development inPhycomyces

Light and temperature markedly influence sporangiophore development inPhycomyces blakesleeanus. Under normal conditions in the dark, low temperature drastically stimulates the production of dwarf sporangiophores (microphorogenesis) and inhibits that of giant sporangiophores (macrophorogenesis). These effects of low temperature could still be observed if applied only for a short period before sporangiophore initiation. Continuous white illumination strongly inhibits microphorogenesis and slightly stimulates macrophorogenesis. Short exposures to white light noticeably inhibit microphorogenesis and stimulate macrophorogenesis when given to mycelia grown for between 90 and 160 h at 14° C or 150 h or more at 10° C. These results indicate the existence in the mycelium of developmental stages for the regulation of sporangiophorogenesis by environmental signals.     

A NEW SPECIES OF SYNCEPHALIS (MUCORALES)

Syncephalis plumigaleata sp. nov. is distinguished from other species by the unilateral placement of the merosporangia on the apical sporangiophore vesicle.     

Positional differentiation for development along the sporangiophore in <Emphasis Type="Italic">Phycomyces blakesleeanus</Emphasis>

 Small segments of Phycomyces sporangiophores regenerate various structures on incubation in a moist chamber. We tested the regeneration capacities of middle sporangiophore segments whose protoplasm had been totally or partially removed and replaced with protoplasm from various segments of a genetically different strain. The structures that were regenerated depended on the source of the injected protoplasm (sporangia of various sizes and segments from middle and basal parts of the sporangiophores), implying a positional differentiation of protoplasm along the sporangiophore axis. Protoplasm from various sources showed a high affinity; that is, they mixed successfully and led, in most cases, to the formation of heterokaryotic regenerating structures. The highest affinity was seen when mixing protoplasm from the middle segments of two different strains. Received: November 29, 2000 / Accepted: March 19, 2002     

PALAEOSTACHYA DIRCEI N. SP., AN AUTHIGENICALLY CEMENTED EQUISETALEAN STROBILUS FROM THE MIDDLE PENNSYLVANIAN OF SOUTHERN ILLINOIS

Equisetalean strobili normally are encountered as disarticulated organs. This condition has necessitated the creation of 12 form genera to accommodate the various morphological architectures and anatomical configurations. Taxonomically useful characteristics, which are discernable in permineralized specimens, are rarely distinguishable in coalified compressions due to their destruction during diagenesis and coalification. Therefore, genera established on the position of sporangiophore-trace origin, such as Schimperia Remy and Remy, are untenable when dealing with coalified compressions. Although the two largest genera of strobili, Calamostachys Schimper and Palaeostachya Weiss, may be synonymous, it is proposed that these form genera be retained when dealing with coalified compression specimens. Each genus provides a particular architectural concept for specimens which may not be assignable to the species level. Calamostachys and Palaeostachya are highly overspeciated genera. It is suggested that characteristics necessary to delimit new species include bract height, degree of bract fusion, disposition of bracts, bract: sporangiophore ratio, number of sporangia per sporangiophore, and sexual status.     

“Apical dominance” in the sporangiophore of the fungus Phycomyces

Summary Control, by the spores, over the elongation and the branching of the sporangiophore is described. If the sporangium is removed from a sporangiophore, the sporangiophore stops growing within a few hours. 6–16h later a branch grows from the defunct growing zone. This branch forms a new sporangium and spores, and then starts to elongate again. The original growing zone can be rescued by replacing the sporangium with another sporangium or a sporangium-sized drop of spores. If the original growing zone is rescued, then it continues growing and the sporangiophore does not form a branch. It can only be rescued, however, within the first 60 min after the original sporangium is removed.This work was submitted in partial fulfillment of the requirements for the Ph. D. degree.     

Development of Sporangiophores of Peronospora parasitica (Pers. ex Fr.) Fr.

The morphology of developing sporangiophores of Peronosporaparasitica from wallflower is described, and morphogenesis maybe divided into the following five stages: the sporangiophoreprimordium, unbranched sporangiophore, branched sporangiophore,spore formation and maturation, and formation of the cross wall.The growth of individual sporangiophores in a humidity chamberwas followed under the microscope, and increase in height andincrease in volume measured. The greatest increase in volumewas during spore formation, when the sporangiophore volume mightquadruple within an hour.     

Coemansia furcata sp. nov. and its distribution in Japan and Taiwan

Coemansia furcata, isolated from forest soils of Japan and Taiwan, is described and illustrated. This species is characterised by its sporangiophore branching and relatively large sporocladia. Its distribution in soils of Japan and Taiwan is discussed. Contributions to Sugadaira Montane Research Center, No. 174.     

Gravitropism in Phycomyces: a role for sedimenting protein crystals and floating lipid globules

To elucidate the graviperception of the unicellular fungus, Phycomycesblakesleeanus, sporangiophores were inspected for intracellular structures which relocate with respect to gravity. Two structures, paracrystalline proteins (so-called octahedral crystals) and an aggregate of lipid globules, were identified which showed redistribution upon reorientation of the sporangiophore. Octahedral crystals occur throughout the sporangiophore, including the apical growing zone, and are localized inside vacuoles in which they reside singly or in clusters of up to 40 loosely associated individuals. Upon a 90° reorientation of sporangiophores, crystal clusters sedimented in approximately 50–200 s from the upper to the lower side, corresponding to a speed of 0.5–2 μm s⁻¹. Stage-4 sporangiophores (with sporangium) of three mutants which lack the crystals displayed anormal kinetics of gravitropism and substantially reduced bending angles in comparison to sporangiophores of the wild type. While horizontally placed wild-type sporangiophores reached the vertical position after 10–12 h, the crystal-lacking mutants bent maximally 40°–50° upward. In stage-1 sporangiophores a conspicuous aggregate of lipid globules is positioned about 50 μm below the apex. The globules floated upwards when the sporangiophore was placed horizontally forming in this way a cap-like aggregate. It is proposed that both the sedimenting protein crystals and the upward-floating globules are involved in gravisensing. Received: 23 March 1999 / Accepted: 27 May 1999     

An Increase in Mechanical Extensibility during the Period of Light-stimulated Growth

The sporangiophore of Phycomyces responds to a temporary increase in light intensity with a transient increase in growth rate that begins 2 to 3 minutes after the initiation of the stimulus and continues until approximately the 12th minute. Tensile tests conducted on the stage IVb sporangiophore demonstrate that an increase in mechanical extensibility of the cell wall occurs 2 minutes after the initiation of a light stimulus and continues until approximately the 15th minute. This finding supports the theory that light-stimulated plant cell expansion and rate of expansion is a function of the mechanical extensibility of the cell wall.     

The Distribution of Cytoplasmic Vesicles, Multivesicular Bodies and Paramural Bodies in Elongating Sporangiophores and Swelling Sporangia of Thamnidium elegans Link

In Thamnidium elegans Link, cytoplasmic vesicles of variablesize were present in large numbers in sporangiophore apicesin agar and in smaller numbers in sporangiophore apices in air.Golgi-like cisternae were associated with small, sub-apicalclusters of vesicles. Apices in agar contained a nucleus andmitochondrion-free tip zone in which vesicles were concentrated.This zone was almost completely absent from apices in air. Inswelling sporangia, vesicles were sparce and were not concentratedagainst the wall. Rates of surface area increase were similarfor sporangiophores elongating, respectively, in agar and inair. Rates of surface area increase during sporangium swellingwere equal to or greater than rates of surface area increaseduring sporangiophore elongation. Vesicles were associated withformation of a secondary wall layer in swollen sporangia. Paramuralbodies and multivesicular bodies were present at all stagesof sporangiophore elongation and sporangium swelling. Isolatedhalo bodies (apical corpuscles) were present in walls at sporangiophoretips, and clusters of similar bodies were present in side wallsof sporangiophores.     

Negative phototropism in the piloboloid mutants of Phycomyces blakesleeanus Bgff.

The sporangiophore (spph) of a piloboloid mutant, genotype pil, of Phycomyces ceases elongation and expands radially in the growth zone shortly after reaching the developmental stage IV b. The pil spph is always negatively phototropic to unilateral visible light when its diameter exceeds 210 m. Photoinduction of spph initiation, light-growth response, threshold of light energy fluence rate for the negative phototropism, avoidance and gravitropism in the pil mutant are all normal. In liquid paraffin, the pil spph shows negative phototropism as does the wild-type spph. Genetic analyses indicate that the negative phototropism of the pil mutant is governed by the phenotypic characteristics of pil but not by specific gene(s) responsible for negative phototropism. These facts imply that the reverse phototropism of the pil mutant results from a loss of the convergent lens effect of the cell because of the increase in cell diameter.Abbreviations spph(s) sporangiophore(s) - wt(s) wild type(s)     

The gravireceptor of Phycomyces. Its development following gravity exposure

The gravitropism of a mature stage IV Phycomyces sporangiophore has a shorter and more uniform latency if the sporangiophore is exposed horizontally to gravity during its earlier development (stage II and stage III). This early exposure to an altered gravitational orientation causes the sporangiophore to develop a gravireceptor as it matures to stage IV and resumes elongation. A technique has been developed to observe the spatial relationship between the vacuole and the protoplasm of a living sporangiophore and to show the reorganization caused by this exposure to altered gravity. Possible gravireceptor mechanisms are discussed.     

Physiological Studies in the Mucorales: PART I THE PHOTOTROPISM OF SPORANGIOPHORES OF PHYCOMYCES BLAKESLEEANUS

Previous views on the physical basis of phototropism in Phycomycessporangiophores are briefly discussed.

1. It was confirmed thatunilaterally illuminated sporangiophoresimmersed in liquidparaffin show strong negative phototropism.
2. Elongation growthceased and no phototropic response took placeunder anaerobicconditions.
3. By focusing a fine beam of light on to one edgeof the growingzone of a sporangiophore, leaving the other sidein darkness,it was established that greater elongation tookplace in theilluminated zone, the sporangiophore tending tobend out ofthe beam. Rapid reversal of the curvature followedwhen theillumination was transferred to the opposite edge ofthe sporangiophore.

Wassink and Boumann's suggestion that phototropism can be initiatedby a one-quantum-per-cell process is criticized in the lightof this result and other work by Castle.