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NASA NTRSDate not supplied

Plant Water Management in Microgravity

The NASA Plant Water Management (PWM) technology demonstrations aboard ISS apply recent advances in microgravity capillary fluidics research towards the mundane yet problematic challenges of simply watering plants in space. Plant growth in a low-g environment is often hampered by inadequate aeration and over-saturation of the root zone. The present effort aims to exploit the passive capillary forces of poorly wetting liquids (i.e., contaminated water) within unique system geometries that effectively replace the role of gravity in providing sufficient aeration and hydration for simulated plants. Several flight demonstrations have been completed on ISS, including soil and hydroponic models in single and parallel channel networks. Two future demonstrations are still in work and plan to further develop the system to handle sustained plant growth with additional sensors to monitor the growth environment. The results to date demonstrate proof-of-concept, system stability, limits of operation, and more, for simulated plant models. Eventually, real plants will be incorporated into these systems and tested on orbit. The implications are discussed in relation to plant growth facilities for further near-term microgravity plant science research as well as for automated food production for long duration human exploration missions.

microgravity↗

NASA NTRSDate not supplied

Plants in Space and Space Crop Production

As astronauts venture farther from Earth, and for longer periods, the space food system will increase in importance. Crop production can supplement a packaged diet to provide additional nutrients and dietary variety for astronauts, and in future missions, bioregenerative approaches may be used to generate a larger percentage of the diet. Plants may also provide behavioural health benefits and assist with other life support functions. Several unique challenges exist for growth of plants in microgravity and on other planetary surfaces like the moon and Mars. Testing with the Veggie and Advanced Plant Habitat (APH) chambers on the International Space Station is allowing us to understand the impacts of gravity and spaceflight on crop growth, nutritional content, acceptability, and the importance of plants to astronauts living and working away from Earth. We are also gaining a better understanding of food safety concerns and the behaviour of space plant microbiomes and plant pathogens. As we transition from research towards operational space crop production to enable human exploration, there are several gaps and challenges of growing crops in space that must be addressed. Research and technology development in key areas such as water and nutrient delivery, plant health monitoring, and crop selection are needed to overcome these challenges. Additionally, there are opportunities for breeding or engineering of custom space crops related to plant growth and development, plant physiology, produce nutrition, organoleptic acceptability, and post-harvest characteristics. Solutions to help ensure food security off-Earth may also translate to better approaches to terrestrial sustainable crop production.

Advanced Plant Habitat↗

NASA NTRS2023

Plants in Space and Space Crop Production

Advanced Plant Habitat↗

NASA NTRS1997

Anatomical features of pepper plants (Capsicum annuum L.) grown under red light-emitting diodes supplemented with blue or far-red light

Pepper plants (Capsicum annuum L. cv., Hungarian Wax) were grown under metal halide (MH) lamps or light-emitting diode (LED) arrays with different spectra to determine the effects of light quality on plant anatomy of leaves and stems. One LED (660) array supplied 90% red light at 660 nm (25nm band-width at half-peak height) and 1% far-red light between 700-800nm. A second LED (660/735) array supplied 83% red light at 660nm and 17% far-red light at 735nm (25nm band-width at half-peak height). A third LED (660/blue) array supplied 98% red light at 660nm, 1% blue light between 350-550nm, and 1% far-red light between 700-800nm. Control plants were grown under broad spectrum metal halide lamps. Plants were gron at a mean photon flux (300-800nm) of 330 micromol m-2 s-1 under a 12 h day-night photoperiod. Significant anatomical changes in stem and leaf morphologies were observed in plants grown under the LED arrays compared to plants grown under the broad-spectrum MH lamp. Cross-sectional areas of pepper stems, thickness of secondary xylem, numbers of intraxylary phloem bundles in the periphery of stem pith tissues, leaf thickness, numbers of choloplasts per palisade mesophyll cell, and thickness of palisade and spongy mesophyll tissues were greatest in peppers grown under MH lamps, intermediate in plants grown under the 660/blue LED array, and lowest in peppers grown under the 660 or 660/735 LED arrays. Most anatomical features of pepper stems and leaves were similar among plants grown under 660 or 660/735 LED arrays. The effects of spectral quality on anatomical changes in stem and leaf tissues of peppers generally correlate to the amount of blue light present in the primary light source.

NASA Discipline Life Support Systems↗

NASA NTRS1995

Growth and photomorphogenesis of pepper plants under red light-emitting diodes with supplemental blue or far-red lighting

Light-emitting diodes (LEDs) are a potential irradiation source for intensive plant culture systems and photobiological research. They have small size, low mass, a long functional life, and narrow spectral output. In this study, we measured the growth and dry matter partitioning of 'Hungarian Wax' pepper (Capsicum annuum L.) plants grown under red LEDs compared with similar plants grown under red LEDs with supplemental blue or far-red radiation or under broad spectrum metal halide (MH) lamps. Additionally, we describe the thermal and spectral characteristics of these sources. The LEDs used in this study had a narrow bandwidth at half peak height (25 nm) and a focused maximum spectral output at 660 nm for the red and 735 nm for the far-red. Near infrared radiation (800 to 3000 nm) was below detection and thermal infrared radiation (3000 to 50,000 nm) was lower in the LEDs compared to the MH source. Although the red to far-red ratio varied considerably, the calculated phytochrome photostationary state (phi) was only slightly different between the radiation sources. Plant biomass was reduced when peppers were grown under red LEDs in the absence of blue wavelengths compared to plants grown under supplemental blue fluorescent lamps or MH lamps. The addition of far-red radiation resulted in taller plants with greater stem mass than red LEDs alone. There were fewer leaves under red or red plus far-red radiation than with lamps producing blue wavelengths. These results indicate that red LEDs may be suitable, in proper combination with other wavelengths of light, for the culture of plants in tightly controlled environments such as space-based plant culture systems.

NASA Discipline Life Support Systems↗

NASA NTRS1981

AgRISTARS: Supporting research. Spring small grains planting date distribution model

A model was developed using 996 planting dates at 51 LANDSAT segments for spring wheat and spring barley in Minnesota, Montana, North Dakota, and South Dakota in 1979. Daily maximum and minimum temperatures and precipitation were obtained from the cooperative weather stations nearest to each segment. The model uses a growing degree day summation modified for daily temperature range to estimate the beginning of planting and uses a soil surface wetness variable to estimate how a fixed number of planting days are distributed after planting begins. For 1979, the model predicts first, median, and last planting dates with root mean square errors of 7.91, 6.61, and 7.09 days, respectively. The model also provides three or four dates to represent periods of planting activity within the planting season. Although the full model was not tested on an independent data set, it may be suitable in areas other than the U.S. Great Plains where spring small grains are planted as soon as soil and air temperatures become warm enough in the spring for plant growth.

Hodges, T.↗

NASA NTRS1982

Increase of cold tolerance in cotton plant (Gossypium hirsutum L.) by mepiquat chloride

Three mepiquat chloride (MC) concentrations - 40, 70, and 100 g a.i./ha - were used to spray cotton (Gossypium hirsutum L., cultival McNair 220) plants to determine whether or not MC would increase their cold tolerance. Seven to ten days after the spray, the plants were exposed to three different cold treatments. No important difference in cold damage was noticed between the control and the MC-treated plants when they were exposed repeatedly to 4.5 C. No plants died when exposed to 0.5 C for 12 h; however, 90% of the 1st and 2nd leaves of the control plants were damaged. This was three times more damage than those leaves of plants treated with 70 and 100 g a.i./ha MC concentrations; 60% f the control and 10-20% of the MC-treated plants died when the plants were subjected to a cold hardening process with 15.5 C day (12 h) and 1.7 C night (12 h) for 10 days, and then, held at -2.2 C for 24 hours. The electrolyte leakage and reflectance measurement data showed that the cell membranes of the MC-treated plants sustained much less damage than those of the control. Freezing injury was easily assessed by reflectance measurements at the 1.65 micrometer wavelength.

Gausman, H. W.↗

NASA NTRS1985

Methane emissions to the atmosphere through aquatic plants

The movement of methane (CH4) from anaerobic sediments through the leaves, stems, and flowers of aquatic plants and into the atmosphere was found to provide a significant pathway for the emission of CH4 from the aquatic substrates of flooded wetlands. Methane concentrations well above the surrounding ambient air levels were found in the mesophyll of 16 varies of aquatic plants and are attributed to transpiration, diffusion, and pressure-induced flow of gaseous CH4 from the roots when they are embedded in CH4-saturated anaerobic sediments. Methane emissions from the emergent parts of aquatic plants were measured using floating chamber techniques and by enclosing the plants in polyethylene bags of known volume. Concentration changes were monitored in the trapped air using syringes and gas chromatographic techniques. Vertical profiles of dissolved CH4 in sediment pore water surrounding the aquatic plants' rhizomes were obtained using an interstitial sampling technique. Methane emissions from the aquatic plants studied varied from 14.8 mg CH4/d to levels too low to be detectable. Rooted and unrooted freshwater aquatic plants were studied as well as saltwater and brackish water plants. Included in the experiment is detailed set of measurements on CH4 emissions from the common cattail (Typha latifolia). This paper illustrates that aquatic plants play an important gas exchange role in the C cycle between wetlands and the atmosphere.

Sebacher, D. I.↗

NASA NTRS1992

A summary of porous tube plant nutrient delivery system investigations from 1985 to 1991

The Controlled Ecological Life Support System (CELSS) Program is a research effort to evaluate biological processes at a one person scale to provide air, water, and food for humans in closed environments for space habitation. This program focuses currently on the use of conventional crop plants and the use of hydroponic systems to grow them. Because conventional hydroponic systems are dependent on gravity to conduct solution flow, they cannot be used in the microgravity of space. Thus, there is a need for a system that will deliver water and nutrients to plant roots under microgravity conditions. The Plant Space Biology Program is interested in investigating the effect that the space environment has on the growth and development of plants. Thus, there is also a need to have a standard nutrient delivery method for growing plants in space for research into plant responses to microgravity. The Porous Tube Plant Nutrient Delivery System (PTPNDS) utilizes a hydrophilic, microporous material to control water and nutrient delivery to plant roots. It has been designed and analyzed to support plant growth independent of gravity and plans are progressing to test it in microgravity. It has been used successfully to grow food crops to maturity in an earth-bound laboratory. This document includes a bibliography and summary reports from the growth trials performed utilizing the PTPNDS.

Dreschel, T. W.↗

NASA NTRS1994

Spectral filtering for plant production

Both plants and animals have one general commonality in their perception of light. They both are sensitive primarily to the 400 to 700 nm wavelength portion of the electromagnetic spectrum. This is referred to as the visible spectrum for animals and as the photosynthetically active radiation (PAR) spectrum for plants. Within this portion of the spectrum, animals perceive colors. Relatively recently it has been learned that within this same spectral range plants also demonstrate varying responses at different wavelengths, somewhat analogous to the definition of various colors at specific wavelengths. Although invisible to the human eye, portions of the electromagnetic spectrum on either side of the visible range are relatively inactive photosynthetically but have been found to influence important biological functions. These portions include the ultraviolet (UV approximately equal to 280-400 nm) and the far-red (FR approximately equal to 700-800 nm). The basic photoreceptor of plants for photosynthesis is chlorophyll. It serves to capture radiant energy which combined with carbon dioxide and water produces oxygen and assimulated carbon, used for the synthesis of cell wall polysaccarides, proteins, membrane lipids and other cellular constituents. The energy and carbon building blocks of photosynthesis sustain growth of plants. On the other hand, however, there are other photoreceptors, or pigments, that function as signal transducers to provide information that controls many physiological and morphological responses of how a plant grows. Known photomorphogenic receptors include phytochrome (the red/far-red sensor in the narrow bands of 655-665 nm and 725-735 nm ranges, respectively) and 'cryptochrome' (the hypothetical UV-B sensor in the 280-320 nm range). Since the USDA team of W. L. Butler, S. B. Hendricks, H. A. Borthwick, H. A. Siegleman and K. Norris in Beltsville, MD detected by spectroscopy, extracted and identified phytochrome as a protein in the 1950's, many other investigators have found evidence of its control functions in plants. Considerably less, however, is known about the yet non-isolated cryptochrome. The information-transferring roles of photoreceptors in plants at specific spectral ranges quite naturally stimulated plant scientists and engineers to consider physically manipulating light to achieve desired physiological and morphological characteristics. One way to manipulate light is to filter it through materials that selectively transmit portions of the sun's spectrum in and near the PAR range.

Young, Roy E.↗

NASA NTRS2002

Development of Biocompatibility Procedures for Assessment of Plant Growth in Ground Test Hardware for the EMCS

The European Modular Cultivation System (EMCS) is an European Space Agency-developed facility designed to support plant research in microgravity on the IS NASA is responsible for providing US specific hardware to use within the EMCS. In preparation for flight, research will be developed and tested at Ames Research Center in the EMCS ground test hardware, the Experiment Reference Module (ERM) In order to determine the acceptability of the ERM for such purposes, biocompatibility tests will be performed to determine that the hardware functions as intended and successfully supports the' growth of plants. In this report, we describe the development of procedures and the collection of baseline data against which to compare ERM function, e.g. biocompatibility testing. A simple and robust system was developed to grow whole Arabidopsis thaliana plants within the confined volumes characteristic of spaceflight hardware. Our system for growing plants eliminated the necessity of a water/nutrient delivery system and allowed for quantifiable assessment of individual plants, as well as entire population dynamics. To insure uniform germination, seeds were started in small straw segments and transplanted into modified scintillation vials. Seedlings were selected prior to transplantation to decrease genetic variability. Plants were grown for a total of 24 days in standard laboratory plant growth chambers under controlled conditions. Sequential digital still images were taken on a daily basis. Analysis of these images allowed for the quantification of even minute environmental effect, on growth dynamics whole plants. The data collected provide reliable growth curves against which to compare plants grown in the ERM.

Bowman, R. N.↗

NASA NTRS2003

Demonstration of a Porous Tube Hydroponic System to Control Plant Moisture and Growth

Accurate remote detection of plant health indicators such as moisture, plant pigment concentrations, photosynthetic flux, and other biochemicals in canopies is a major goal in plant research. Influencing factors include complex interactions between wavelength dependent absorbing and scattering features from backgrounds as well as canopy biochemical and biophysical constituents. Accurately controlling these factors in outdoor field studies is difficult. Early testing of a porous tube plant culture system has indicated that plant biomass production, biomass partitioning, and leaf moisture of plants can be controlled by precisely managing the root water potential. Managing nutrient solution chemistry can also control plant pigments, biochemical concentrations, plant biomass production, and photosynthetic rates. A test bed was developed which utilized the porous tube technology with the intent of evaluating remote sensing systems, spectral analyses procedures, gas-exchange, and fluorescence measurements for their ability to detect small differences in plant water status. Spectral analysis was able to detect small differences in the mean leaf water content between the treatments. However these small differences were not detectable in the gas-exchange or fluorescence measurements.

Dreschel, T. W.↗

NASA NTRS2010

Root Zone Respiration on Hydroponically Grown Wheat Plant Systems

Root respiration is a biological phenomenon that controls plant growth and physiological development during a plant's lifespan. This process is dependent on the availability of oxygen in the system where the plant is located. In hydroponic systems, where plants are submerged in a solution containing vital nutrients but no type of soil, the availability of oxygen arises from the dissolved oxygen concentration in the solution. This oxygen concentration is dependent on the , gas-liquid interface formed on the upper surface of the liquid, as given by Henry's Law, depending on pressure and temperature conditions. Respiration rates of the plants rise as biomass and root zone increase with age. The respiration rate of Apogee wheat plants (Triticum aestivum) was measured as a function of light intensity (catalytic for photosynthesis) and CO2 concentration to determine their effect on respiration rates. To determine their effects on respiration rate and plant growth microbial communities were introduced into the system, by Innoculum. Surfactants were introduced, simulating gray-water usage in space, as another factor to determine their effect on chemical oxygen demand of microbials and on respiration rates of the plants. It is expected to see small effects from changes in CO2 concentration or light levels, and to see root respiration decrease in an exponential manner with plant age and microbial activity.

Soler-Crespo, R. A.↗

NASA NTRS2003

Device for Automated Cutting and Transfer of Plant Shoots

A device that enables the automated cutting and transfer of plant shoots is undergoing development for use in the propagation of plants in a nursery or laboratory. At present, it is standard practice for a human technician to use a knife and forceps to cut, separate, and grasp a plant shoot. The great advantage offered by the present device is that its design and operation are simpler than would be those of a device based on the manual cutting/separation/grasping procedure. [The present device should not be confused with a prior device developed for partly the same purpose and described in Compliant Gripper for a Robotic Manipulator (NPO-21104), NASA Tech Briefs, Vol. 27, No. 3 (March 2003), page 59.]. The device (see figure) includes a circular tube sharpened at its open (lower) end and mounted on a robotic manipulator at its closed (upper) end. The robotic manipulator simply pushes the sharpened open end of the tube down onto a bed of plants and rotates a few degrees clockwise then counterclockwise about the vertical axis, causing the tube to cut a cylindrical plug of plant material. Exploiting the natural friction between the tube and plug, the tube retains the plug, without need for a gripping mechanism and control. The robotic manipulator then retracts the tube, translates it to a new location over a plant-growth tray, and inserts the tube part way into the growth medium at this location in the tray. A short burst of compressed air is admitted to the upper end of the tube to eject the plug of plant material and drive it into the growth medium. A prototype has been tested and verified to function substantially as intended. It is projected that in the fully developed robotic plant-propagation system, the robot control system would include a machine- vision subsystem that would automatically guide the robotic manipulator in choosing the positions from which to cut plugs of plant material. Planned further development efforts also include more testing and refinement of the design and operation described above.

Cipra, Raymond↗

NASA NTRSDate not supplied

System Modeling of a Lunar Molten Regolith Electrolysis Plant

Introduction: In-Situ Resource Utilization (ISRU) is the process of extracting local resources to produce commodities for propulsion, life support systems, and off-planet construction rather than transporting consumables from Earth. Molten Regolith Electrolysis (MRE) is a novel ISRU method of extracting oxygen gas and metal alloy from lunar regolith. The MRE process involves placing lunar regolith between two electrodes, through which current is passed, to melt the regolith and reduce the metal oxide constituents by direct electrolysis (e.g. FeO, SiO2, MgO, Al2O3) into oxygen gas and metal alloys. The oxygen is liquefied and used as propellant for landers, while the metals (e.g. Ferro-alloys) are further processed and used in structural building materials and parts manufacturing. A system model was developed that accounted for the major processes of an MRE plant (from excavation of raw materials to storage of products) to assess the feasibility of a lunar MRE plant. The System Engineering and Integration (SE&I) ISRU Modeling and Analysis (SIMA) team utilized its previously documented system sizing model, the Mission Analysis and Integration Tool (MAIT) [1] as framework of the system model. MAIT uses MATLAB/Simulink to integrate subsystem models into a complete system model of the MRE plant. Total mass, volume, and power requirements were computed for numerous iterations of a MRE plant. System Model: Figure 1: MRE Plant Block Diagram The regolith excavation model determines the mass and power needed to excavate sufficient regolith. The preheating auger initiates the regolith heating process before regolith enters the MRE re-actor to reduce the energy required to turn the solid into a molten liquid. The MRE reactor is modeled in COMSOL Multiphysics and based on the research by Dominguez, Sibille, and Schreiner [2, 3, 4]. This preliminary reactor model provides an accurate calculation of thermal equilibrium during electrochemical operation of the reactor system to assess the optimal mass and power required to process the inlet flow of regolith. The model also computes the outlet flowrates of oxygen and molten products. For this analysis, the primary components of the metal alloy considered were iron and silicon. The oxygen is then purified using an Yttrium Stabilized Zirconia (YSZ) electrode, followed by liquefaction using a 90K cryocooler to be stored as liquid oxygen in insulated cylindrical tanks. In future iterations of the system model, the molten metal tapped from the MRE reactor will undergo additional processing or refinement. However, downstream handling of metals is currently a technology gap that is missing a high TRL subsystem model. Therefore, for this analysis, the accumulated metal alloy stream terminates after leaving the MRE reactor. Study Goals: This analysis investigates multiple input variables to the system to determine the sensitivity of a (near) complete plant at full-scale. This preliminary investigation ran parametric sweeps on the MRE reactor geometry, electrical current supply, layers of multi-layer insulation (MLI) on the reactor, size of the electrodes in the oxygen purification model, and regolith composition (based on landing site location). Three production targets of oxygen (1,000, 10,000, and 50,000 kg/yr) were investigated for this analysis. The parametric sweeps conducted in this analysis provide valuable insight into the expected impact of the various model inputs on plant size. This information can be used to identify the most critical components of the plant and guide future decisions on allocating funding for research and development, providing subsystem developers with appropriate interfaces with downstream and upstream processes, and assessing the overall feasibility of MRE when compared to other ISRU plants. References: [1] Carlson, A. et al. (2024) ICES, ICES-2024-53. [2] Dominguez, D.A., and Sibille, L. (2011) AIAA, AIAA-2011-700. [3] Schreiner, S.S. (2015) MIT, Dissertation. [4] Schreiner, S.S. et al. (2016) ASR, 57(7), pp.1585-1603.

ISRU↗

NASA NTRS2000

How can plants tell which way is up?

Many people think of plants as essentially sessile organisms that do not actively respond to their environment. What could be further from the truth! In fact, plants are capable of a variety of movements, including the dramatic nastic responses (such as Venus fly trap closure) and the less sensational tropisms. These latter movements are directed growth responses to some type of external stimulus such as gravity (gravitropism, formerly known as geotropism) or light (phototropism). This paper describes some interesting exercises that are derived from recent work, including research that has led to experiments performed on two Space Shuttle missions in 1997 (Kiss et al. 1998). The study of tropisms can be a useful way to introduce students to plant biology in high school and introductory college courses. In our experience, students are fascinated by plant movements when they are presented in lectures and find laboratory experiences on this topic quite engaging. Laboratory work on plant tropisms can also be used to introduce important concepts in science such as hypothesis testing, quantitative analysis, and the use of statistics. The laboratory exercises described in this paper involve the higher plant Arabidopsis thaliana, which has become an important organism in molecular biology research and is the focus of an international plant genome project. Based on the material presented here, a number of plant gravitropism laboratory exercises with Arabidopsis that are simple in terms of equipment/materials and procedures can be developed. These exercises are robust in that they work well even in the hands of introductory students, and they can be expanded according to the individual instructor's needs. This paper describes two exercises that have been performed by beginning college students, and these exercises can easily be performed in biology classes in most high school settings.

Non-NASA Center↗

NASA NTRS1997

Spectral quality affects disease development of three pathogens on hydroponically grown plants

Plants were grown under light-emitting diode (LED) arrays with various spectra to determine the effects of light quality on the development of diseases caused by tomato mosaic virus (ToMV) on pepper (Capsicum annuum L.), powdery mildew [Sphaerotheca fuliginea (Schlectend:Fr.) Pollaci] on cucumber (Cucumis sativus L.), and bacterial wilt (Pseudomonas solanacearum Smith) on tomato (Lycopersicon esculentum Mill.). One LED (660) array supplied 99% red light at 660 nm (25 nm bandwidth at half-peak height) and 1% far-red light between 700 to 800 nm. A second LED (660/735) array supplied 83% red light at 660 nm and 17% far-red light at 735 nm (25 nm bandwidth at half-peak height). A third LED (660/BF) array supplied 98% red light at 660 nm, 1% blue light (BF) between 350 to 550 nm, and 1% far-red light between 700 to 800 nm. Control plants were grown under broad-spectrum metal halide (MH) lamps. Plants were grown at a mean photon flux (300 to 800 nm) of 330 micromoles m-2 s-1 under a 12-h day/night photoperiod. Spectral quality affected each pathosystem differently. In the ToMV/pepper pathosystem, disease symptoms developed slower and were less severe in plants grown under light sources that contained blue and UV-A wavelengths (MH and 660/BF treatments) compared to plants grown under light sources that lacked blue and UV-A wavelengths (660 and 660/735 LED arrays). In contrast, the number of colonies per leaf was highest and the mean colony diameters of S. fuliginea on cucumber plants were largest on leaves grown under the MH lamp (highest amount of blue and UV-A light) and least on leaves grown under the 660 LED array (no blue or UV-A light). The addition of far-red irradiation to the primary light source in the 660/735 LED array increased the colony counts per leaf in the S. fuliginea/cucumber pathosystem compared to the red-only (660) LED array. In the P. solanacearum/tomato pathosystem, disease symptoms were less severe in plants grown under the 660 LED array, but the effects of spectral quality on disease development when other wavelengths were included in the light source (MH-, 660/BF-, and 660/735-grown plants) were equivocal. These results demonstrate that spectral quality may be useful as a component of an integrated pest management program for future space-based controlled ecological life support systems.

NASA Center KSC↗

NASA NTRS1993

Calcium and signal transduction in plants

Environmental and hormonal signals control diverse physiological processes in plants. The mechanisms by which plant cells perceive and transduce these signals are poorly understood. Understanding biochemical and molecular events involved in signal transduction pathways has become one of the most active areas of plant research. Research during the last 15 years has established that Ca2+ acts as a messenger in transducing external signals. The evidence in support of Ca2+ as a messenger is unequivocal and fulfills all the requirements of a messenger. The role of Ca2+ becomes even more important because it is the only messenger known so far in plants. Since our last review on the Ca2+ messenger system in 1987, there has been tremendous progress in elucidating various aspects of Ca(2+) -signaling pathways in plants. These include demonstration of signal-induced changes in cytosolic Ca2+, calmodulin and calmodulin-like proteins, identification of different Ca2+ channels, characterization of Ca(2+) -dependent protein kinases (CDPKs) both at the biochemical and molecular levels, evidence for the presence of calmodulin-dependent protein kinases, and increased evidence in support of the role of inositol phospholipids in the Ca(2+) -signaling system. Despite the progress in Ca2+ research in plants, it is still in its infancy and much more needs to be done to understand the precise mechanisms by which Ca2+ regulates a wide variety of physiological processes. The purpose of this review is to summarize some of these recent developments in Ca2+ research as it relates to signal transduction in plants.

NASA Program Space Biology↗