[The growth movements of moss protonemata under clinostatic and microgravity conditions]. / Rostovye dvizheniia protonemy mkhov v usloviiakh klinostatirovaniia i mikrogravitatsii.

Populations of dark-grown protonemata of moss Ceratodon purpureus wt-4 (Germany) and wt-U (Ukraine) were rotated on clinostat or flown in space (experiment "Protonema" aboard Bion-11, December 24, 1996-January 7, 1997) to determine the effects of altered gravity on orientation of protonemata growing filaments. Protonemata had been cultivated 8 days in vertical stationary position at dark to be transported to microgravity or placed in clinostat for the period of 14 days. In the ground control, protonemata demonstrated the negatively gravitropic growth (straight upwards in a bundle of compact filaments). The horizontal or circular rotation in clinostat and exposure to microgravity made filaments grow every each way within the substrate plane but with an apparent trend to rightward curling resulting in "spiral galaxies".

The gravireaction of Ceratodon protonemata treated with gibberellic acid.

Moss protonemata exhibit negative gravitropism and the amyloplasts of the apical cell seem to play a key role in protonemal gravisensitivity. However, the mechanisms of this process are still poorly understood. Previously, we have shown that Ceratodon protonemata grown on agar-medium demonstrated greater gravicurvature than protonemata grown on medium with 11 mM glucose. In this study, we have examined whether gibberellic acid (GA), which promotes alpha-amylase expression, influences graviresponse of C. purpureus protonemata (strains WT-4 and WT-U) and how this event interacts with exogenous soluble sugars. After gravistimulation the WT-4 strain curved about twice as fast as the WT-U strain. However, responses of both strains to added substances were similar. High concentration of glucose (0.11 M) caused a decrease in protonema curvature, while the same concentration of sucrose did not significantly change the angles of curvature compared with controls. GA at 0.1 mM and higher concentrations inhibited gravitropism, and caused some apical cells to swell. The possible involvement of the carbohydrates in gravitropism is discussed.

Gravitropism and phototropism in protonemata of the moss Pohlia nutans (Hedw.) Lindb.

The gravitropism of protonemata of Pohlia nutans is described and compared with that of other mosses. In darkness, protonemata showed negative gravitropism. Under uniform illumination they grew radially over the substrate surface, whereas unilateral illumination induced positive phototropic growth. Gravitropism was coupled with starch synthesis and amyloplast formation. Protonematal gravitropic growth is more variable than the strict negative gravitropism of Ceratodon chloronema.

Influence of gravity on spatial orientation and morphogenesis of moss sporophytes.

During the growth and development of the sporophytic capsules of some moss species, negative gravitropism is changed for a positive one. Horizontal clinostat rotation induced unregulated growth of the sporophytes and their twisting; some of sporophytes remained straight, however. It has been established that the change of the gravitropic reaction is related to capsule formation and to the redistribution of amyloplast cells of the sporophyte graviperception zone.

Gravi- and photostimuli in moss protonema growth movements.

Moss protonemal growth direction is controlled by at least three factors, photo-, gravi- and autotropism. It is possible to experimentally separate these factors and to control selectively their morphological appearance. In darkness protonema grow negatively gravitropically, and unilateral illumination initiated positive phototropism. Red light suppressed auto- and gravitropism, blue light suppressed only gravitropism. Green light allowed both gravi- and autotropism. The effect of light on gravitropism might involve changes in starch synthesis.

Gravitropism in caulonemata of the moss Pottia intermedia.

The gravitropism of caulonemata of Pottia intermedia is described and compared with that of other mosses. Spore germination produces primary protonemata including caulonemata which give rise to buds that form the leafy moss plant, the gametophore. Primary caulonemata are negatively gravitropic but their growth and the number of filaments are limited in the dark. Axenic culture of gametophores results in the production of secondary caulonemata that usually arise near the leaf base. Secondary protonemata that form in the light are agravitropic. Secondary caulonemata that form when gametophores are placed in the dark for several days show strong negative gravitropism and grow well in the dark. When upright caulonemata are reorientated to the horizontal or are inverted, upward bending can be detected after 1 h and caulonemata reach the vertical within 1-2 d. Clear amyloplast sedimentation occurs 10-15 minutes after horizontal placement and before the start of upward curvature. This sedimentation takes place in a sub-apical zone. Amyloplast sedimentation also takes place along the length of upright and inverted Pottia protonemata. These results support the hypothesis that amyloplast sedimentation functions in gravitropic sensing since sedimentation occurs before gravitropism in Pottia and since the location and presence of a unique sedimentation zone is conserved in all four mosses known to gravitropic protonomata.

Growth of Pottia intermedia protonemata in altered gravity.

Plants are immobile; therefore, they are oriented in space due to growth movements--tropisms. The latter occur in response to environmental stimuli such as gravity (gravitropism), light (phototropism), chemical compounds or water (chemo- and hydrotropisms). Gravity is the only force that was impossible to control. The moss protonemata are among the limited group of plant objects with tip growth. What is unique about this structure is that protonemal apical cells both sense and respond to gravity. It is considered that the apical cell perceives gravity through amyloplasts (Sack, 1993; Chaban, 1996). Although the dynamics of protonemata negative gravitropism in different moss species was studied in detail, the role of gravity in both the structural polarity of apical cells and the formation of protonematal mat with circular symmetry is completely unexplored. Using the unique possibility to fly the moss on the space shuttle (STS-87) we aimed in this study to analyze the character of the interaction of gravity with light and endogenous factors in the pattern of protonemata space orientation.

Behavior of amyloplasts in photo- and gravitropism of the moss protonema.

The protonema of mosses Ceratodon purpureus and Pottia intermedia is negatively gravitropic in darkness and grows on the substrate surface under illumination. However, the putative mechanisms of these growth responses are not well understood so far. For gravitropism, sedimentation of amyloplasts has been widely assumed to be the first step of the signal transduction chain. This model was supported by numerous observations where amyloplasts' number or size in a protonema apical cell correlated with its gravisensitivity. Unlike multicellular graviperceptive organs, a protonema apical cell is the same site for both of gravity and light perception and realization of growth movements. In addition, red light is known to change the cell responses to gravity. Therefore, we analysed the influence of red light on the events associated with graviperception and growth movements of protonema apical cells, namely: plastid behavior, size, and number, starch content, chlorophyll fluorescence intensity and alpha-amylase activity, under gravistimulation of dark-grown protonema.

Calcium gradient in plant cells with polarized growth in simulated microgravity.

Plant cells characterized by apical growth, for example, root hairs and apical cells of moss protonema, are a convenient model to address the problem of gravity response mechanisms including initiation of cell polarity. The fluorescent calcium probe, chlorotetracycline, allowed us to display the calcium distribution gradient in these cells. Irradiation by red light led to a sharp decrease in the Ca2+ ion activity in cells. During clinostatting in darkness the pattern of calcium influx and distribution changes inconsiderably as compared with control; in root hairs calcium is detected mainly in their apices and bases as in control. Addition of chlorpromazine to the medium probably increases the influx and accumulation of Ca2+ ions. Under data obtained confirm speculations on the Ca2+ ion functional role for the apical growth of plant cells and may suggest the participation of gravity in redistribution or activation of ion channels, calcium channels included, in the plasmalemma.

[Polarity of cell permeability and its control by phytochrome]. / Poliarnost' kletochnoi pronitsaemosti i ee kontrol' fitokhromom.

Growing chloronema and rhizoid cells of the intact protonema of Funaria hygrometrica absorb luminescent stains selectively. The cation stains--acridine orange and neutral red, at pH 5.0 are absorbed by growing tips of the apical cells. The anion stain (uranine) is absorbed by the basal end of the apical cells and by intercalary cells of the protonema filaments. Red light inactivates the cell permeability to the cation and anion stains. Far-red light partly reversed the action of the red light solely on the initial stages of the irradiation. During the red light induced branching of apical cells the cation stains enter the cells through a new originated growing point.