Characterization of Adaptation in Phototropism of Arabidopsis thaliana

Phototropic curvature has been measured for etiolated Arabidopsis thaliana seedlings with and without a preirradiation. A bilateral preirradiation with 450-nm light at a fluence greater than about 0.1 micromole per square meter causes a rapid desensitization to a subsequent 450-nanometer unilateral irradiation at 0.5 micromole per square meter. Following a refractory period, the capacity to respond phototropically recovers to the predesensitization level, and the response is then enhanced. The length of the refractory period is between 10 and 20 minutes. Both the time needed for recovery and the extent of enhancement increase with increasing fluence of the bilateral preirradiation. Based on the relative spectral sensitivities of desensitization and enhancement, these responses can be separated. Desensitization is induced by blue light but not by red light. Enhancement, however, is induced by both blue and red light. Thus, enhancement can be induced without desensitization but not vice versa. Both desensitization and enhancement affect only the magnitude of the response and do not affect the fluence threshold.     

Growth Distribution during Phototropism of Arabidopsis thaliana Seedlings

The elongation rates of two opposite sides of hypocotyls of Arabidopsis thaliana seedlings were measured during phototropism by using an infrared imaging system. In first positive phototropism, second positive phototropism, and red light-enhanced first positive phototropism, curvature toward the light source was the result of an increase in the rate of elongation of the shaded side and a decrease in the rate of elongation of the lighted side of the seedlings. The phase of straightening that followed maximum curvature resulted from a decrease in the elongation rate of the shaded side and an increase in the elongation rate of the lighted side. These data for the three types of blue light-induced phototropism tested in this study and for the phase of straightening are all clearly consistent with the growth rate changes predicted by the Cholodny-Went theory.     

Blue and Green Light-Induced Phototropism in Arabidopsis thaliana and Lactuca sativa L. Seedlings

Exposure time-response curves for blue and green light-induced phototropic bending in hypocotyls of Arabidopsis thaliana (L.) Heynh. and Lactuca sativa L. seedlings are presented. These seedlings show significant phototropic sensitivity up to 540 to 550 nanometers. Since wave-lengths longer than 560 nanometers do not induce phototropic bending, it is suggested that the response to 510 to 550 nanometers light is mediated by the specific blue light photoreceptor of phototropism. We advise care in the use of green `safelights' for studies of phototropism.     

A Common Fluence Threshold for First Positive and Second Positive Phototropism in Arabidopsis thaliana

The relationship between the amount of light and the amount of response for any photobiological process can be based on the number of incident quanta per unit time (fluence rate-response) or on the number of incident quanta during a given period of irradiation (fluence-response). Fluence-response and fluence rate-response relationships have been measured for second positive phototropism by seedlings of Arabidopsis thaliana. The fluence-response relationships exhibit a single limiting threshold at about 0.01 micromole per square meter when measured at fluence rates from 2.4 × 10⁻⁵ to 6.5 × 10⁻³ micromoles per square meter per second. The threshold values in the fluence rateresponse curves decrease with increasing time of irradiation, but show a common fluence threshold at about 0.01 micromole per square meter. These thresholds are the same as the threshold of about 0.01 micromole per square meter measured for first positive phototropism. Based on these data, it is suggested that second positive curvature has a threshold in time of about 10 minutes. Moreover, if the times of irradiation exceed the time threshold, there is a single limiting fluence threshold at about 0.01 micromole per square meter. Thus, the limiting fluence threshold for second positive phototropism is the same as the fluence threshold for first positive phototropism. Based on these data, we suggest that this common fluence threshold for first positive and second positive phototropism is set by a single photoreceptor pigment system.     

Mutants of Arabidopsis thaliana with altered responses to auxins and gravity

Auxin-resistant mutants of Arabidopsis have been induced and isolated by screening for survivors on a medium containing the herbicide 2,4-D. Thirty independently arisen mutants have been isolated in this way and one of them, P 83, has been investigated in detail. When wild type and P 83 are compared in concentration/response curves, where the response is the inhibition of root growth, the ED₅₀ values of the auxins, 2,4-D and IAA, are 14-fold higher for the mutant. The mutant also responds differently to gravity: its roots do not show positive geotropism, but tend to grow with a clockwise curvature on agar surfaces. The seedling roots of the mutant also grow more rapidly than those of the wild type in the absence of 2,4-D, following faster germination. The F₁ between P 83 and wild type is similar to the latter, but has a slightly increased resistance to 2,4-D. Results obtained from the F₂, F₃ and backcross generations suggest monofactorial inheritance. Most of the other 29 mutants have the P 83 phenotype, but at least five are different. Four have lower levels of resistance to 2,4-D and P 83, and their roots appear to respond normally to gravity. One mutant has an abnormal georesponse and a much higher level of resistance to 2,4-D than P 83.     

Effect of simulated microgravity on auxin polar transport in inflorescence axis of Arabidopsis thaliana.

The morphology, growth and development of higher plants are strongly influenced by environmental stimuli on the earth, which affect the changes in the dynamics of plant hormones in plants. Qualitative and quantitative changes in plant hormones are the most important internal factor to regulate plant growth and development. Among them, auxin (IAA) is of most significant. There are numerous reports concerning the physiological roles of auxin in plant growth and development (Matthysse and Scott 1984). One of the characteristics of auxin is to have the ability of polar transport along the vector of gravity on the earth (Schneider and Wightman 1978), suggesting that the activity of auxin polar transport is also important for the growth and development of plants. It has recently been reported that the normal activity of auxin polar transport in inflorescence axis of Arabidopsis thaliana was required for flower formation (Okada et al. 1991, Ueda et al. 1992). Considering the above evidence together with the fact that gravity affects the morphology, growth and development of higher plants, gravity might affect the qualitative and quantitative changes in plant hormones including the activity of auxin polar transport. In this paper, we report the effect of microgravity condition simulated by a three-dimensional (3-D) or a horizontal clinostat on the activity of auxin polar transport in inflorescence axis of Arabidopsis thaliana.     

Phototropism and gravitropism in lateral roots of Arabidopsis

Gravitropism and, to a lesser extent, phototropism have been characterized in primary roots, but little is known about structural/functional aspects of these tropisms in lateral roots. Therefore, in this study, we report on tropistic responses in lateral roots of Arabidopsis thaliana. Lateral roots initially are plagiogravitropic, but when they reach a length of approximately 10 mm, these roots grow downward and exhibit positive orthogravitropism. Light and electron microscopic studies demonstrate a correlation between positive gravitropism and development of columella cells with large, sedimented amyloplasts in wild-type plants. Lateral roots display negative phototropism in response to white and blue light and positive phototropism in response to red light. As is the case with primary roots, the photoresponse is weak relative to the graviresponse, but phototropism is readily apparent in starchless mutant plants, which are impaired in gravitropism. To our knowledge, this is the first report of phototropism of lateral roots in any plant species.     

Touch modulates gravity sensing to regulate the growth of primary roots of Arabidopsis thaliana

Plants must sense and respond to diverse stimuli to optimize the architecture of their root system for water and nutrient scavenging and anchorage. We have therefore analyzed how information from two of these stimuli, touch and gravity, are integrated to direct root growth. In Arabidopsis thaliana, touch stimulation provided by a glass barrier placed across the direction of growth caused the root to form a step-like growth habit with bends forming in the central and later the distal elongation zones. This response led to the main root axis growing parallel to, but not touching the obstacle, whilst the root cap maintained contact with the barrier. Removal of the graviperceptive columella cells of the root cap using laser ablation reduced the bending response of the distal elongation zone. Similarly, although the roots of the gravisensing impaired pgm1-1 mutant grew along the barrier at the same average angle as wild-type, this angle became more variable with time. These observations imply a constant gravitropic re-setting of the root tip response to touch stimulation from the barrier. In wild-type plants, transient touch stimulation of root cap cells, but not other regions of the root, inhibited both subsequent gravitropic growth and amyloplast sedimentation in the columella. Taken together, these results suggest that the cells of the root cap sense touch stimuli and their subsequent signaling acts on the columella cells to modulate their graviresponse. This interaction of touch and gravity signaling would then direct root growth to avoid obstacles in the soil while generally maintaining downward growth.     

Microbial antibiotic production aboard the International Space Station

Previous studies examining metabolic characteristics of bacterial cultures have mostly suggested that reduced gravity is advantageous for microbial growth. As a consequence, the question of whether space flight would similarly enhance secondary metabolite production was raised. Results from three prior space shuttle experiments indicated that antibiotic production was stimulated in space for two different microbial systems, albeit under suboptimal growth conditions. The goal of this latest experiment was to determine whether the enhanced productivity would also occur with better growth conditions and over longer durations of weightlessness. Microbial antibiotic production was examined onboard the International Space Station during the 72-day 8A increment. Findings of increased productivity of actinomycin D by Streptomyces plicatus in space corroborated with previous findings for the early sample points (days 8 and 12); however, the flight production levels were lower than the matched ground control samples for the remainder of the mission. The overall goal of this research program is to elucidate the specific mechanisms responsible for the initial stimulation of productivity in space and translate this knowledge into methods for improving efficiency of commercial production facilities on Earth.     

The development of the International Space Station centrifuge.

Gravitational biology research facility "Centrifuge" is currently under development for the International Space Station. Research in the Complex Organism Biology, indispensable to the progress in Health Science, is only possible in the Centrifuge aboard the station. So, on-orbit 1 G controls for various specimens including small mammals, fish, and higher plants will be rigorously done in the Centrifuge. This facility is also capable of providing "reduced gravity" likely on the Moon or on Mars. Thus, it will play a key role in creating knowledge of space fundamental biology. As part of the offset of NASA's Shuttle launch services for the Japanese Experiment Module, JAXA is developing the Centrifuge Rotor (CR), the Life Sciences Glovebox (LSG) and the Centrifuge Accommodation Module (CAM). Critical Design Review (CDR) of LSG was conducted on July 2004, while the system CDRs of the CAM and CR are scheduled for December 2004 and August 2005, respectively. Their launch schedules are under review.