Photoreactions in Phycomyces : Responses to the stimulation of narrow test areas with ultraviolet light

Equipment has been developed for ultraviolet illumination of sharply bounded test areas of the growing zone of sporangiophores of Phycomyces. The growing zone is opaque for this light and the tropic responses are negative. Periodic short narrow stimuli on alternating sides produce periodic tropic responses when applied at x > 0.5 mm, but none for x < 0.5 mm, where x is the distance below the sporangium. Sustained tropic stimuli, applied at constant x, produce tropic responses for any x > 0.1 mm. The lag between stimulus and response is 3.3 min. for any x > 0.5 mm. For smaller x the lag increases progressively. In all cases the tropic bend occurs at values of x > 0.5 mm. Sustained tropic stimuli, applied at constant height relative to ground, produce relatively sharp tropic bends. The center of the bend is at all times close to the simultaneous position of the stimulated area. The boundaries of a light-adapted zone move less than 0.1 mm in 10 min. relative to the sporangium. It is concluded that the receiving and adapting structures do not move relative to the sporangium, and that the responding system does not move relative to ground. The two systems move relative to each other with the speed of growth. The responding system does not extend above x = 0.5 mm.     

ORIENTATION AND LOCUS OF TROPIC PHOTORECEPTOR MOLECULES IN SPORES OF BOTRYTIS AND OSMUNDA

Study of the tropic responses of Botrytis cinerea and Osmunda cinnamomea spores to blue light shows the photoreceptor molecules to be highly dichroic and oriented: in Botrytis their axes of maximum absorption lie perpendicular to the nearby cell surface; in Osmunda, parallel. The chief evidence lies in a comparison of their responses to plane polarized light—both germinate parallel to the vibration planes (defined by the axis of vibration of the electric vector and the axis of light propagation)—with those to partial illumination with unpolarized light: Botrytis grows from its brighter part; Osmunda, from its darker. The degree of orientation produced by polarized light corresponds, at high intensities, to that produced by the imposition of such large (about 100 per cent) intensity differences across a cell as to preclude all alternatives to oriented dichroic receptors. The photoreceptors of the Botrytis spore lie within the cell wall's inner half. The chief evidence lies in the component of its tropic responses to polarized light within the vibration plane: germination peaks about 10° off the vibration axis. This deviation arises from focusing which is effective only in the wall's inner half. At high intensities, anomalies appear in Botrytis which are interpreted as "centering," i.e., a tendency toward growth from the center of two or more equally illuminated points of a cell rather than from one of them.     

Physical strain-mediated microtubule reorientation in the epidermis of gravitropically or phototropically stimulated maize coleoptiles

During gravitropic and phototropic curvature of the maize coleoptile, the cortical microtubules (MTs) adjacent to the outer epidermal cell wall assume opposite orientations at the two sides of the organ. Starting from a uniformly random pattern during straight growth in darkness, the MTs reorientate perpendicularly to the organ axis at the outer (faster growing) side and parallel to the organ axis at the inner (slower growing) side. As similar reorientations can be induced during straight growth by increasing or decreasing the effective auxin concentration, it has been proposed that these reorientations may be used as a diagnostic test for assessing the auxin status of the epidermal cells during tropic curvature. This idea was tested by determining the MT orientations in the coleoptile of intact maize seedlings in which the gravitropic or phototropic curvature was prevented or inversed by an appropriate mechanical counterforce. Forces that just prevented the coleoptile from curving in a gravity or light field prevented reorientations of the MTs. Forces strong enough to overcompensate the tropic stimuli by enforcing curvature in the opposite direction induced reorientations of the MTs opposite to those produced by tropic stimulation. These results show that the MTs at the outer surface of the coleoptile respond to changes in mechanical tissue strain rather than to gravitropic or phototropic stimuli and associated changes at the level of auxin or any other element in the signal transduction chain between perception of tropic stimuli and asymmetric growth response. It is proposed that cortical MTs can act as strain gauges in a positive feed-back regulatory circle utilized for amplification and stabilization of environmentally induced changes in the direction of elongation growth.     

Relationships between height on the shore and size distributions of Thais SPP. (Gastropoda: Muricidae)

Average body sizes in a variety of marine animals have been reported to be correlated with height in the intertidal zone. This paper reports field observations on four species of Thais showing that the relationship between size distribution and intertidal height or subtidal depth is highly variable. Intertidal size gradients in Thais have been explained by a size dependent set of responses to light and gravity, but experiments reported here do not indicate that snails of different sizes behave differently with respect to the intertidal gradient per se and it is argued that, when there is a size gradient, it is produced by the direct responses of snails to factors such as shelter, temperature, desiccation, and food. It is suggested that a response to token stimuli is unlikely to evolve when the token stimuli are poorly correlated with the relevant environmental factors, especially if the latter can be monitored directly by the animal.     

Blue light sensory systems in plants

Plant development is influenced by many environmental stimuli, including light, temperature and gravity. Of these stimuli, light is of particular importance because plants depend on it for energy and, thus, for survival. Moreover, virtually all stages of plant development are regulated in part by light through the action of various photosensory systems. Examples of light-regulated processes include germination, stem growth, leaf and root development, tropic responses and flower induction. This review provides an analysis of recent investigations of blue light sensory systems in plants. Current results suggest that plants respond to blue light through a complex photosensory network that incorporates the action of multiple blue light perception systems.     

Tropic Responses of Phycomyces Sporangiophores to Gravitational and Centrifugal Stimuli

A low-speed centrifuge was used to study the tropic responses of Phycomyces sporangiophores in darkness to the stimulus of combined gravitational and centrifugal forces. If this stimulus is constant the response is a relatively slow tropic reaction, which persists for up to 12 hours. The response is accelerated by increasing the magnitude of the gravitational-centrifugal force. A wholly different tropic response, the transient response, is elicited by an abrupt change in the gravitational-centrifugal stimulus. The transient response has a duration of only about 6 min. but is characterized by a high bending speed (about 5°/min.). An analysis of the distribution of the transient response along the growing zone shows that the active phase of the response has a distribution similar to that of the light sensitivity for the light-growth and phototropic responses. Experiments in which sporangiophores are centrifuged in an inert dense fluid indicate that the sensory mechanism of the transient response is closely related to the physical deformation of the growing zone caused by the action of the gravitational-centrifugal force on the sporangiophore as a whole. However, the response to a steady gravitational-centrifugal force is most likely not connected with this deformation, but is probably triggered by the shifting of regions or particles of differing density relative to one another inside the cell.     

A hormonal regulatory module that provides flexibility to tropic responses

Plants orient their growth depending on directional stimuli such as light and gravity, in a process known as tropic response. Tropisms result from asymmetrical accumulation of auxin across the responding organ relative to the direction of the stimulus, which causes differential growth rates on both sides of the organ. Here, we show that gibberellins (GAs) attenuate the gravitropic reorientation of stimulated hypocotyls of dark-grown Arabidopsis (Arabidopsis thaliana) seedlings. We show that the modulation occurs through induction of the expression of the negative regulator of auxin signaling INDOLE-3-ACETIC ACID INDUCIBLE19/MASSUGU2. The biological significance of this regulatory mechanism involving GAs and auxin seems to be the maintenance of a high degree of flexibility in tropic responses. This notion is further supported by observations that GA-deficient seedlings showed a much lower variance in the response to gravity compared to wild-type seedlings and that the attenuation of gravitropism by GAs resulted in an increased phototropic response. This suggests that the interplay between auxin and GAs may be particularly important for plant orientation under competing tropic stimuli.     

A molecular basis for the physiological variation in shade avoidance responses: A tale of two ecotypes

Using two ecotypes of Stellaria longipes with contrasting responses to shade, we found that plants can differ in their responses to similar light cues, reflecting adaptations to their natural habitat. It was also observed that the plants could distinguish between distinct shade signals. Furthermore, the activity of wall modifying proteins, expansins and xyloglucan endotransglucosylase/hydrolase(s) (XTHs) was regulated during these responses. However, only expansin activity and gene expression profiles correlated with observed growth trends. The differential expression of expansins was light signal specific and ecotype specific and could account for both the trends in growth and their magnitude. We have thus established a potential molecular basis for the observed plasticity in responses to shade.Key words: shade avoidance, cell wall modification, expansins, XTHs, Stellaria longipes, phenotypic plasticity, light quality     

The Light Growth Response of Phycomyces

With the help of an automated tracking system we have studied the characteristics of the transient light growth response of Phycomyces. The response shows a sharply defined latency. The Q₁₀ of the reciprocal latency is 2.4. Response patterns at different peaks of the action spectrum are the same. The gradual variation of response magnitude over a wide range of adapted intensifies parallels that of phototropism. The responses to saturating stimuli exhibit a strong oscillation with a constant period of 1.6 min and variable damping. The growth responses to sinusoidally varying light intensities show a system bandwidth of 2.5 x 10^-3 Hz. The linear dependence of phase shift on frequency is largely attributable to the latency observed with pulse stimuli. In the high intensity range a previously suspected increase of the steady-state growth rate with intensity has been confirmed. The light growth responses of mutants selected for diminished phototropism have been investigated. Many of these mutants have sizable but grossly distorted growth responses.     

Phototropism and polarotropism of primary chloronemata of the moss Physcomitrella patens: responses of mutant strains

The phototropic and polarotropic responses of primary chloronemata grown from germinated minated spores of three mutant strains of the moss, Physcomitrella patens, have been studied and compared with those of the wild-type. The mutants and wild-type show the same qualitative tropic responses but differ with respect to the light conditions under which they are expressed. In both the wild-type and mutants the responses are controlled by phytochrome. In monochromatic red light, at low fluence rates, wild-type primary chloronemata grow positively phototropically in unidirectional light or perpendicular to the electrical vector (E) in polarised light; at high fluence rates growth in unidirectional light is lateral to the incident light or, in polarised light, parallel to E. The mutants, however, show only the lateral phototropic or parallel polarotropic responses at all fluence rates of red light tested. In far-red light, the wild-type primary chloronemata adopt a positive phototropic or a perpendicular polarotropic response; the mutants show the same responses but in a lower percentage of filaments. These results and those at other wavelengths indicate either that the mutants are impaired in their ability to adopt the positive phototropic and perpendicular polarotropic responses or that in the mutants the transition between the “low light” (positive phototropic-perpendicular polarotropic) and the “high light” (lateral phototropic-parallel polarotropic) responses is shifted to a lower photon fluence rate. Possible explanations of this phenotypic difference are discussed.     

Plant tropisms: providing the power of movement to a sessile organism

In an attempt to compensate for their sessile nature, plants have developed growth responses to deal with the copious and rapid changes in their environment. These responses are known as tropisms and they are marked by a directional growth response that is the result of differential cellular growth and development in response to an external stimulation such as light, gravity or touch. While the mechanics of tropic growth and subsequent development have been the topic of debate for more than a hundred years, only recently have researchers been able to make strides in understanding how plants perceive and respond to tropic stimulations, thanks in large part to mutant analysis and recent advances in genomics. This paper focuses on the recent advances in four of the best-understood tropic responses and how each affects plant growth and development: phototropism, gravitropism, thigmotropism and hydrotropism. While progress has been made in deciphering the events between tropic stimulation signal perception and each characteristic growth response, there are many areas that remain unclear, some of which will be discussed herein. As has become evident, each tropic response pathway exhibits distinguishing characteristics. However, these pathways of tropic perception and response also have overlapping components - a fact that is certainly related to the necessity for pathway integration given the ever-changing environment that surrounds every plant.     

The regulation of cell wall extensibility during shade avoidance: a study using two contrasting ecotypes of Stellaria longipes

Shade avoidance in plants involves rapid shoot elongation to grow toward the light. Cell wall-modifying mechanisms are vital regulatory points for control of these elongation responses. Two protein families involved in cell wall modification are expansins and xyloglucan endotransglucosylase/hydrolases. We used an alpine and a prairie ecotype of Stellaria longipes differing in their response to shade to study the regulation of cell wall extensibility in response to low red to far-red ratio (R/FR), an early neighbor detection signal, and dense canopy shade (green shade: low R/FR, blue, and total light intensity). Alpine plants were nonresponsive to low R/FR, while prairie plants elongated rapidly. These responses reflect adaptation to the dense vegetation of the prairie habitat, unlike the alpine plants, which almost never encounter shade. Under green shade, both ecotypes rapidly elongate, showing that alpine plants can react only to a deep shade treatment. Xyloglucan endotransglucosylase/hydrolase activity was strongly regulated by green shade and low blue light conditions but not by low R/FR. Expansin activity, expressed as acid-induced extension, correlated with growth responses to all light changes. Expansin genes cloned from the internodes of the two ecotypes showed differential regulation in response to the light manipulations. This regulation was ecotype and light signal specific and correlated with the growth responses. Our results imply that elongation responses to shade require the regulation of cell wall extensibility via the control of expansin gene expression. Ecotypic differences demonstrate how responses to environmental stimuli are differently regulated to survive a particular habitat.     

Mechanical properties of muscle spindles in Xenopus laevis

Mechanical properties of isolated living muscle spindles from Xenopus laevis were examined in order to determine their role in sensory transduction. The reticular zone of the intrafusal muscle fibers was identified microscopically by: (1) its position beneath the sensory endings, (2) its length, 50–100 μm, (3) its extension during intrafusal muscle contraction, and (4) its coarse striations with a period of about 1.5 times the normal sarcomere length. The reticular zone in the passive muscle spindle did not extend until the spindle was stretched to about 1.05–1.1 its maximal length in the animal (L _m ). Evidence was obtained that the absence of extension of the reticular zone at normal muscle lengths was due to the presence of the spindle capsule which acted as a stiff element in parallel with the sensory region. At those lengths at which the reticular zone did extend (> L _m ), no rate — sensitive mechanical properties were detected in response to step and ramp extensions. The sensory discharge of the spindle showed no dynamic transient in response to ramp extensions if the reticular zone were not extended. During extension of the reticular zone a dynamic sensory transient appeared. It is concluded that current notions on the mechanical origin of the rate — sensitive properties of the sensory discharge of the muscle spindle do not apply to Xenopus laevis. In addition, it is not likely that the passive spindle in this animal is a sensitive stretch receptor.     

Rice plants response to the disruption of OsCSLD4 gene

The plant cell wall is a complex polysaccharide network and performs important developmental and physiological functions far beyond supplying the physical constrains. Plant cells have the ability to react to cell wall defects as exhibited by changes in gene expression, accumulation of ectopic lignin, stress responses and growth arrest. It is a major challenge to understand how plants sense and respond to wall integrity since very little is known about the signaling involved in the responses. Cellulose synthase-like D (CSLD) proteins mediating the biosynthesis of a wall polysaccharide polymer make up a common subfamily to all plants. Recently, we have reported the functional characterization of CSLD4 in rice. Mutations in OsCSLD4 show morphological alterations and pleiotropic effects on wall compositions and structure. Our study demonstrates that OsCSLD4 play a critical role in cell wall formation and plant growth. Here we show the subtle wall alterations by separating the culm residues into five fractions. Quantitative RT-PCR analysis further revealed that the expression of various genes involved in xylan synthesis and cell cycle regulation was altered in mutant plants, as the responses to OsCSLD4 disruption. Therefore, plants may have fine sensory machinery to react to wall defects and modulate growth for adapting to the changes.Key words: OsCSLD4, cell wall biosynthesis, plant development, wall integrity, rice     

Crosslinked elastic fibers are necessary for low energy loss in the ascending aorta

In the large arteries, it is believed that elastin provides the resistance to stretch at low pressure, while collagen provides the resistance to stretch at high pressure. It is also thought that elastin is responsible for the low energy loss observed with cyclic loading. These tenets are supported through experiments that alter component amounts through protease digestion, vessel remodeling, normal growth, or in different artery types. Genetic engineering provides the opportunity to revisit these tenets through the loss of expression of specific wall components. We used newborn mice lacking elastin (Eln^−/−) or two key proteins (lysyl oxidase, Lox^−/−, or fibulin-4, Fbln4^−/−) that are necessary for the assembly of mechanically-functional elastic fibers to investigate the contributions of elastic fibers to large artery mechanics. We determined component content and organization and quantified the nonlinear and viscoelastic mechanical behavior of Eln^−/−, Lox^−/−, and Fbln4^−/− ascending aorta and their respective controls. We confirmed that the lack of elastin, fibulin-4, or lysyl oxidase leads to absent or highly fragmented elastic fibers in the aortic wall and a 56–97% decrease in crosslinked elastin amounts. We found that the resistance to stretch at low pressure is decreased only in Eln^−/− aorta, confirming the role of elastin in the nonlinear mechanical behavior of the aortic wall. Dissipated energy with cyclic loading and unloading is increased 53–387% in Eln^−/−, Lox^−/−, and Fbln4^−/− aorta, indicating that not only elastin, but properly assembled and crosslinked elastic fibers, are necessary for low energy loss in the aorta.     

Avoidance response, house response, and wind responses of the sporangiophore of Phycomyces

Avoidance response: An object placed 1 mm from the growing zone of a Phycomyces sporangiophore elicits a tropic response away from the object. The dependence of this response on the size of the object and its distance from the specimen is described, as well as measurements which exclude electric fields, electromagnetic radiation, temperature, and humidity as avoidance-mediating signals. This response is independent of the composition and surface properties of the object and of ambient light. House Response: A house of 0.5- to 10-cm diameter put over a sporangiophore elicits a transient growth response. Avoidance responses inside closed houses are slightly smaller than those in the open. Wind responses: A transverse wind elicits a tropic response into the wind, increasing with wind speed. A longitudinal wind, up or down, elicits a transient negative growth response to a step-up in wind speed, and vice versa. It is proposed that all of the effects listed involve wind sensing. This proposal is supported by measurements of aerodynamic effects of barriers and houses on random winds. The wind sensing is discussed in terms of the hypothesis that a gas is emitted by the growing zone (not water or any normal constituent of air), the concentration of which is modified by the winds and monitored by a chemical sensor. This model puts severe constraints on the physical properties of the gas.     

Cell wall extension behavior of Phycomyces sporangiophores during the pressure response.

The cylindrical, single-celled sporangiophore of Phycomyces blakesleeanus grows (enlarges) predominantly in the longitudinal direction during two stages of development; stage I and stage IVb. Cell enlargement (cell wall extension) occurs in a distinct region termed the "growing zone." It was previously reported that a large step-up or pulse-up in turgor pressure, greater than approximately 0.02 MPa, will elicit a transient decrease in longitudinal growth rate of the stage I and stage IVb sporangiophore. This transient decrease in longitudinal growth rate is termed the "pressure response." Both the magnitude and duration of the pressure response depend on the magnitude of the turgor pressure step-up or pulse-up. Qualitatively, the pressure response is similar to the stretch response, which is produced with the application of a longitudinal force (load) on the sporangiophore. In this investigation, the growth (extension) behavior of the cell wall in the growing zone is studied during the pressure response. It is found that both the extension rate of the cell wall in the growing zone and the length of the growing zone decrease during the pressure response, and that together they account for the observed decrease in longitudinal growth rate.     

Mechanosensing and thigmomorphogenesis,a physiological and biomechanical point of view

Plant sensitivity to mechanical stimuli is obvious when observing the movements of Mimosa pudica leaflets when they are touched [1] or those of the Venus fly trap [2]. It is now well established that other plants are also sensitive to mechanical stimuli even if they do not exhibit such rapid movements [1]. There is a renewal of interest in mechanical stimuli as very important cues for the control of plant growth [3] and morphogenesis [4], [5]. This review focuses on mechanosensing in the case of external mechanical loading and its effect on the growth patterns of plant organs (thigmomorphogenesis). The first part of this paper deals with the responses at the whole plant level and their ecological significance. The second part deals with the perception process, with emphasis on the variable that is perceived by the plant. Knowledge about mechanosensors is not presented in great detail because this area of intensive research has been recently reviewed [6], [7]. The third part focuses on transduction, i.e., early responses at the cellular level, and particularly focuses on the importance of the kinetics of loading and the kinetics of cellular responses for the interpretation of experimental results. The fourth part focuses on parameters that regulate the mechanosensing process and points out the importance of quantitative studies. Because thigmomorphogenesis and gravitropism are difficult to disentangle, the review ends with data on gravitropism where mechanosensing is involved.     

Tropic Responses of Funaria Spores to Red Light

The tropic responses of Funaria hygrometrica spores to continuous illumination with red light (610 to 690 mμ) have been studied over the intensity range from 10^-5 through 10⁺⁶ erg/cm² second, using both plane polarized light and partial illumination with unpolarized light. From the relative frequency of outgrowth origin in different directions, the following is inferred. (1). The germination direction of chloronemal filaments is directly influenced by red light over this whole intensity range, while that of rhizoids tends to be opposite the chloronema. (2) Three photoreceptor systems direct chloronemal primordia: (a) A low intensity system acting from 10^-5 to 10^-0.5 erg/cm² second. It favors their growth from a cell's brightest part(s). Its photoreceptors are disoriented, excited by the electric vector, and probably are dispersed phytochrome molecules. (b) A medium intensity system which acts largely alone only at 10^0.5 erg/cm² second but is influential from 10⁰ to 10⁵ erg/cm² second. It likewise favors growth from a cell's brightest part(s); its receptor molecules are also excited electrically, but they are tangentially oriented. (c) A high intensity system which acts alone from 10⁵ to 10⁶ erg/cm² second and is influential down to 10¹ erg/cm² second. It favors growth of the chloronemas from a cell's darkest part. Its receptors probably are magnetically excited and tangentially oriented. The polarotropic responses of the chloronemas resemble those directing their origins. One new feature is that under intense (10⁶ erg/cm² second) plane polarized and vertically directed light, many soon grow to form tight helices.