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Plant photomorphogenesis and canopy growth

An important motivation for studying photomorphogenesis is to understand the relationships among plant photophysiology in canopies, canopy productivity, and agronomic yield. This understanding is essential to optimize lighting systems used for plant farming in controlled environments (CE) and for the design of genetically engineered crop strains with altered photoresponses. This article provides an overview of some basic principles of plant photomorphogenesis in canopies and discusses their implications for (1) scaling up information on plant photophysiology from individual plants in CE to whole canopies in the field, and (2) designing lighting conditions to increase plant productivity in CE used for agronomic purposes (e.g. space farming in CE Life Support Systems). We concentrate on the visible (lambda between 400 and 700 nm) and far-infrared (FR; lambda greater than 700 nm) spectral regions, since the ultraviolet (UV; 280 to 400 nm) is covered by other authors in this volume.

Ballare, Carlos L.↗

Photomorphogenesis, photosynthesis, and seed yield of wheat plants grown under red light-emitting diodes (LEDs) with and without supplemental blue lighting

Red light-emitting diodes (LEDs) are a potential light source for growing plants in spaceflight systems because of their safety, small mass and volume, wavelength specificity, and longevity. Despite these attractive features, red LEDs must satisfy requirements for plant photosynthesis and photomorphogenesis for successful growth and seed yield. To determine the influence of gallium aluminium arsenide (GaAlAs) red LEDs on wheat photomorphogenesis, photosynthesis, and seed yield, wheat (Triticum aestivum L., cv. 'USU-Super Dwarf') plants were grown under red LEDs and compared to plants grown under daylight fluorescent (white) lamps and red LEDs supplemented with either 1% or 10% blue light from blue fluorescent (BF) lamps. Compared to white light-grown plants, wheat grown under red LEDs alone demonstrated less main culm development during vegetative growth through preanthesis, while showing a longer flag leaf at 40 DAP and greater main culm length at final harvest (70 DAP). As supplemental BF light was increased with red LEDs, shoot dry matter and net leaf photosynthesis rate increased. At final harvest, wheat grown under red LEDs alone displayed fewer subtillers and a lower seed yield compared to plants grown under white light. Wheat grown under red LEDs+10% BF light had comparable shoot dry matter accumulation and seed yield relative to wheat grown under white light. These results indicate that wheat can complete its life cycle under red LEDs alone, but larger plants and greater amounts of seed are produced in the presence of red LEDs supplemented with a quantity of blue light.

NASA Discipline Life Support Systems↗

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↗

Altering the axial light gradient affects photomorphogenesis in emerging seedlings of Zea mays L

The axial (longitudinal) red light gradient (632 nanometers) of 4 day old dark-grown maize seedlings is increased by staining the peripheral cells of the coleoptile. The magnitude of increase in the light gradient is dependent solely on the light-absorbing qualities of the stain used. Metanil yellow has no effect on the axial red-light gradient, while methylene blue causes a large increase in this light gradient. These stains did not affect growth in darkness or the sensitivity of mesocotyl elongation to red light. However, mesocotyl elongation was altered for the dark-grown seedlings stained with methylene blue when these seedlings were transplanted, covered with soil, and permitted to emerge under natural lighting conditions. These observations are consistent with the idea that there is a single perceptive site below the coleoptilar node, and suggest that this perceptive site gives the actinic light which has traveled downward through the length of the shoot from an entry point in the plant tip region.

NASA Discipline Number 40-20↗

Photomorphogenesis and photoassimilation in soybean and sorghum grown under broad spectrum or blue-deficient light sources

The role of blue light in plant growth and development was investigated in soybean (Glycine max [L.] Merr. cv Williams) and sorghum (Sorghum bicolor [L.] Moench. cv Rio) grown under equal photosynthetic photon fluxes (approximately 500 micromoles per square meter per second) from broad spectrum daylight fluorescent or blue-deficient, narrow-band (589 nanometers) low pressure sodium (LPS) lamps. Between 14 and 18 days after sowing, it was possible to relate adaptations in photosynthesis and leaf growth to dry matter accumulation. Soybean development under LPS light was similar in several respects to that of shaded plants, consistent with an important role for blue light photoreceptors in regulation of growth response to irradiance. Thus, soybeans from LPS conditions partitioned relatively more growth to leaves and maintained higher average leaf area ratios (mean LAR) that compensated lower net assimilation rates (mean NAR). Relative growth rates were therefore comparable to plants from daylight fluorescent lamps. Reductions in mean NAR were matched by lower rates of net photosynthesis (A) on an area basis in the major photosynthetic source (first trifoliolate) leaf. Lower A in soybean resulted from reduced leaf dry matter per unit leaf area, but lower A under LPS conditions in sorghum correlated with leaf chlorosis and reduced total nitrogen (not observed in soybean). In spite of a lower A, mean NAR was larger in sorghum from LPS conditions, resulting in significantly greater relative growth rates (mean LAR was approximately equal for both light conditions). Leaf starch accumulation rate was higher for both species and starch content at the end of the dark period was elevated two- and three-fold for sorghum and soybean, respectively, under LPS conditions. Possible relations between starch accumulation, leaf export, and plant growth in response to spectral quality were considered.

NASA Discipline Life Support Systems↗

Application of sunlight and lamps for plant irradiation in space bases

The radiation sources used for plant growth on a space base must meet the biological requirements for photosynthesis and photomorphogenesis. In addition, the sources must be energy and volume efficient, while maintaining the required irradiance levels, spectral, spatial and temporal distribution. These requirements are not easily met, but as the biological and mission requirements are better defined, then specific facility designs can begin to accommodate both the biological requirements and the physical limitations of a space-based plant growth system.

Sager, J. C.↗

Development of Gravity Sensitive Plant Cells (Ceratodon) in Microgravity

Protonemata of the moss Ceratodon are tip-growing cells that grow up in the dark. This cell type is unique compared to cells in almost any other organism, since the growth of the plant cell itself is completely oriented by gravity. Thus, both the processes of gravity sensing and the gravity response occur in the same cell. Gravity sensing appears to rely upon amyloplasts (starch-filled plastids) that sediment. This sedimentation occurs in specific zones and plastid zonation is complex with respect to plastid morphology, distribution, and gravity. Microtubules restrict the extent of plastid sedimentation (i.e., they are load-bearing). Light also is important since apical cells have a phytochrome-based positive phototropism, light quality influences plastid zonation and sedimentation (photomorphogenesis), and red light suppresses gravitropism at higher but not lower light intensities. Many of these processes were examined in a 16 day spaceflight experiment, "SPM-A" space moss" or "SPAM)) on STS-87 that landed in December, 1997. The work described here involves the definition of a second flight experiment that builds upon the data and questions arising from STS-87. Effort was directed towards further definition of an experiment for the Shuttle (dubbed "SOS" for "Son of SPAM"). Our current target is STS 107 that is scheduled to fly in January 2001. This definition addressed two goals of the STS107 experiment. The goals of the current experiment were to determine whether the cytoskeleton plays a role in maintaining and generating an apical (non-random) plastid distribution in microgravity and to determine the development and extent of clockwise spiral tip-growth in microgravity.

Sack, Fred D.↗

Ion channels and the transduction of light signals

Studies of biological light-sensing mechanisms are revealing important roles for ion channels. Photosensory transduction in plants is no exception. In this article, the evidence that ion channels perform such signal-transducing functions in the complex array of mechanisms that bring about plant photomorphogenesis will be reviewed and discussed. The examples selected for discussion range from light-gradient detection in unicellular algae to the photocontrol of stem growth in Arabidopsis. Also included is some discussion of the technical aspects of studies that combine electrophysiology and photobiology.

NASA Discipline Plant Biology↗

Light-regulated protein and mRNA synthesis in root caps of maize

Illumination of maize roots initiates changes in mRNA levels and in the activities of proteins within the root cap. Using Northern analysis we showed a 5-6 fold increase in the levels of three specific mRNAs and a 14-fold increase in plastid mRNA. This increase is rapid, occurring within 30 minutes of illumination. With prolonged periods of darkness following illumination, messages return to levels observed in dark, control caps. For two species of mRNA illumination results in a reduction in message levels. Light-stimulated increases in the levels of specific mRNAs are proportionally greater than are increases in the activities of corresponding proteins. We suggest that the light-stimulated increase in protein activity in root caps may be preceded by and occur as a consequence of enhanced levels of mRNA. Our work suggests that photomorphogenesis in roots could involve changes in the levels of a wide variety of mRNAs within the root cap.

NASA Program Space Biology↗

Radiation in controlled environments: influence of lamp type and filter material

Radiation in controlled environments was characterized using fluorescent and various high-intensity-discharge (HID) lamps, including metal halide, low-pressure sodium, and high-pressure sodium as the radiation source. The effects of water, glass, or Plexiglas filters on radiation were determined. Photosynthetic photon flux (PPF, 400 to 700 nm), spectra (400 to 1000 nm), shortwave radiation (285-2800 nm), and total radiation (300 to 100,000 nm) were measured, and photosynthetically active radiation (PAR, 400 to 700 nm) and longwave radiation (2800 to 100,000 nm) were calculated. Measurement of PPF alone was not an adequate characterization of the radiation environment. Total radiant flux varied among lamp types at equal PPF. HID lamps provided a lower percentage of longwave radiation than fluorescent lamps, but, when HID lamps provided PPF levels greater than that possible with fluorescent lamps, the amount of longwave radiation was high. Water was the most effective longwave radiation filter. Glass and Plexiglas similarly filtered longwave more than shortwave radiation, but transmission of nonphotosynthetic shortwave radiation was less with Plexiglas than glass. The filter materials tested would not be expected to influence photomorphogenesis because radiation in the action spectrum of phytochrome was not altered, but this may not be the only pigment involved.

NASA Discipline Life Support Systems↗

Characterization of nucleoside triphosphatase activity in isolated pea nuclei and its photoreversible regulation by light

A nucleoside triphosphatase (NTPase) present in highly purified preparations of pea nuclei was partially characterized. The activity of this enzyme was stimulated by divalent cations (Mg2+ = Mn2+ > Ca2+), but was not affected by the monovalent cations, Na+ and K+. The Mg(2+)-dependent activity was further stimulated by concentrations of Ca2+ in the low micromolar range. It could catalyze the hydrolysis of ATP, GTP, UTP, and CTP, all with a pH optimum of 7.5. The nuclear NTPase activity was not inhibited by vanadate, oligomycin, or nitrate, but was inhibited by relatively low concentrations of quercetin and the calmodulin inhibitor, compound 48/80. The NTPase was stimulated more than 50% by red light, and this effect was reversed by subsequent irradiation with far-red light. The photoreversibility of the stimulation indicated that the photoreceptor for this response was phytochrome, an important regulator of photomorphogenesis and gene expression in plants.

NASA Discipline Plant Biology↗

Irradiance and spectral distribution control system for controlled environment chambers

This paper describes a closed-loop control system for controlling the irradiance and spectral quality generated by fluorescent lamps in a controlled environment chamber. The 400 to 800 nm irradiance and the ratio of the red waveband (600 to 700 nm) to the far-red waveband (700 to 800 nm) were independently controlled and varied as functions of time. A suggested application is to investigate the possibility of synergistic effects of changing irradiance levels and changing spectral distributions on photoperiodism and photomorphogenesis.

NASA Center KSC↗