Super Dwarf Research Articles

Bread Wheat in Space Flight: Is There a Difference in Kernel Quality?

Planning long-term space flights necessarily includes issues of providing food for the crew. One of the areas of research is the development of technologies for independent production of food by the crew. Extensive research on lettuce has confirmed that the "space production" of lettuce is not inferior to that on Earth, even in the absence of gravity, but the same deep understanding of the quality of grain crops has not yet been achieved. Therefore, the goal of our work is to establish whether the conditions for growing wheat in outer space without gravity affect the weight and basic parameters of the grain, and whether this leads to increased asymmetry of the kernel and distortion of the starch composition. The objects of the study were wheat (Triticum aestivum L.) kernels of the Super Dwarf cultivar. Of which, 100 kernels matured in outer space conditions in the Lada growth chamber on the International Space Station (ISS), and 85 kernels of the control wheat grown in a similar growth chamber under terrestrial conditions. It has been established that kernels from ISS have significant differences to a smaller extent in weight, area, length, and width of the kernel. However, the kernels under both conditions were predominantly large (the average weight of a kernel in space is 0.0362 g, and in terrestrial conditions-0.0376 g). The hypothesis that the level of fluctuating asymmetry will increase in outer space was not confirmed; significant differences between the options were not proven. In general, the kernels are fairly even (coefficients of variation for the main parameters of the kernel are within 6-12%) and with a low or very low level of asymmetry. The length of starch granules of type A in filled and puny kernels is significantly greater in kernels from ISS than in the control, and in terms of the width of starch granules B and roundness indices, both experimental variants are the same. It can be assumed that the baking qualities of earthly kernels will be slightly higher, since the ratio of type B starch granules to type A is 5-8% higher than on the ISS. Also, the width of the aleurone layer cells in mature kernels was significantly inferior to the result obtained on Earth. The work proposes a new method for establishing the asymmetry of kernels without a traumatic effect (in early works, it was supposed to study asymmetry in transverse sections of the kernels). Perhaps this will make it possible to further develop a computer scanning program that will determine the level of asymmetry of the wheat fruit.

Genome-Wide Association Study Identifies a Plant-Height-Associated Gene OsPG3 in a Population of Commercial Rice Varieties.

Plant height is one of the most crucial components of plant structure. However, due to its complexity, the genetic architecture of rice plant height has not been fully elucidated. In this study, we performed a genome-wide association study (GWAS) to determine rice plant height using 178 commercial rice varieties and identified 37 loci associated with rice plant height (LAPH). Among these loci, in LAPH2, we identified a polygalacturonase gene, OsPG3, which was genetically and functionally associated with rice plant height. The rice plant exhibits a super dwarf phenotype when the knockout of the OsPG3 gene occurs via CRISPR-Cas9 gene-editing technology. RNA-Seq analysis indicated that OsPG3 modulates the expression of genes involved in phytohormone metabolism and cell-wall-biosynthesis pathways. Our findings suggest that OsPG3 plays a vital role in controlling rice plant height by regulating cell wall biosynthesis. Given that rice architecture is one of the most critical phenotypes in rice breeding, OsPG3 has potential in rice's molecular design breeding toward an ideal plant height.

Growth and Photosynthesis Responses of a Super Dwarf Rice Genotype to Shade and Nitrogen Supply

Specific aspects of plant cultivation require tests under fully controlled environmental conditions with restricted energy supply, such as orbit-based space laboratories and low-light conditions. For these growing conditions, super dwarf plants have been developed as model crops, and a gibberellin- deficient Super Dwarf Rice genotype was proposed as a model crop for space flight plant experiments. We tested this genotype in a climate chamber experiment under different illumination and nitrogen supply levels to assess its suitability under scenarios with limited resource availability. A 25% reduction in illumination led to a 75% reduction in yield, mainly due to a 60% reduction in formed tillers and 20% reduction in grain weight, and a 80% reduction in illumination caused total yield loss. Leaf area under reduced illumination was significantly lower, and only marginal changes in the dimensions of leaves were observed. Plant photosynthesis was not significantly different between control and 75% illumination. This was explained by a higher photochemical efficiency under lower light conditions and a reduced mesophyll resistance. Therefore, we concluded that this genotype is well-suited for plant experiments under space and light-limited conditions since it kept its small stature and showed no shade avoidance mechanisms, such as leaf elongation, which would complicate experiments under low-light conditions. Nitrogen concentrations of 2.8 and 1.4 mmol/L led to no differences in plant growth. We concluded that a nitrogen concentration of 1.4 mmol/L is sufficient for this genotype under the light intensities.

Developmental, nutritional and hormonal anomalies of weightlessness-grown wheat

The behavior of water in weightlessness, as occurs in orbiting spacecraft, presents multiple challenges for plant growth. Soils remain saturated, impeding aeration, and leaf surfaces remain wet, impeding gas exchange. Herein we report developmental and biochemical anomalies of “Super Dwarf” wheat (Triticum aestivum L.) grown aboard Space Station Mir during the 1996–97 “Greenhouse 2” experiment. Leaves of Mir-grown wheat were hyperhydric, senesced precociously and accumulated aromatic and branched-chain amino acids typical of tissues experiencing oxidative stress. The highest levels of stress-specific amino acids occurred in precociously-senescing leaves. Our results suggest that the leaf ventilation system of the Svet Greenhouse failed to remove sufficient boundary layer water, thus leading to poor gas exchange and onset of oxidative stress. As oxidative stress in plants has been observed in recent space-flight experiments, we recommend that percentage water content in apoplast free-spaces of leaves be used to evaluate leaf ventilation effectiveness. Mir-grown plants also tillered excessively. Crowns and culms of these plants contained low levels of abscisic acid but high levels of cytokinins. High ethylene levels may have suppressed abscisic acid synthesis, thus permitting cytokinins to accumulate and tillering to occur.

Screening of a broad range of rice (Oryza sativa L.) germplasm for in vitro rapid plant regeneration and development of an early prediction system

Rice has emerged as a model monocot for studies in agriculture and biotechnology due to its relatively small genome and a ready accessibility to plant material. Tissue culture is one of the tools required for genetic transformation and some breeding programs, and the selection of high-frequency regenerator types is essential for success in these technologies. Thirty-three rice entries with agricultural and biotechnological characteristics of interest were screened with the aim to identify the best regenerators. Entries that exhibited between 50% and 90% regeneration frequencies include ‘Taipei-309,’ ‘Super Dwarf,’ ‘Norin’ (japonica types), PI 312777, ‘Ali Combo’ (indica types), ‘STG-S,’ and ‘LA3’ (red rice types). One third of the entries tested were at least two times better at regeneration than the often-cited regenerator ‘Nipponbare.’ Those entries showing at least 85% frequency of greening or somatic embryo formation at 15 or 30 d on regeneration medium ultimately produced whole plants after 45 d on regeneration medium at high frequency (at least 40%); those entries not reaching the 85% threshold of greening by Days 15 or 30 exhibited moderate (15–40%) to low (less than 10%) frequency of whole plant regeneration. This greening response suggests the means for an early prediction system for identification of useful rice regenerator lines, which would be beneficial for high-throughput screening of germplasm as well as for decreasing the time and cost of in vitro culture.

A dwarf wheat mutant is associated with increased drought resistance and altered responses to gravity

Drought resistance is an important trait for crops. Here we report a new wheat mutant with increased drought resistance and altered gravitropism. During the bombardment transformation of Chinese wheat cultivar ‘Jingdong 6’, a recessive remarkably dwarf wheat mutant named ‘s-dwarf’ (super dwarf wheat) was created. It was severely dwarf and gibberellic acid insensitive. The relative water loss ratio of the detached leaf and the transpiration rate of s-dwarf seedlings were remarkably lower than Jingdong 6 while the photosynthetic rate of s-dwarf seedlings was significantly higher than Jingdong 6. Most of the s-dwarf seedlings survived in recovering experiement after water loss. The stalk of s-dwarf seedling also showed reduced gravitropism. This is the first report about a new dwarf wheat mutant associated with increased drought resistance and altered stalk gravitropism.

A dwarf wheat mutant is associated with increased drought resistance and altered responses to gravity

Drought resistance is an important trait for crops. Here we report a new wheat mutant with increased drought resistance and altered gravitropism. During the bombardment transformation of Chinese wheat cultivar ‘Jingdong 6’, a recessive remarkably dwarf wheat mutant named ‘s-dwarf’ (super dwarf wheat) was created. It was severely dwarf and gibberellic acid insensitive. The relative water loss ratio of the detached leaf and the transpiration rate of s-dwarf seedlings were remarkably lower than Jingdong 6 while the photosynthetic rate of s-dwarf seedlings was significantly higher than Jingdong 6. Most of the s-dwarf seedlings survived in recovering experiement after water loss. The stalk of s-dwarf seedling also showed reduced gravitropism. This is the first report about a new dwarf wheat mutant associated with increased drought resistance and altered stalk gravitropism.

Characterizing the Environmental Response of a Gibberellic Acid–Deficient Rice for Use as a Model Crop

Rice (Oryza sativa L.) is a useful model crop plant. Rice was the first crop plant to have its complete genome sequenced. Unfortunately, even semidwarf rice cultivars are 60 to 90 cm tall, and large plant populations cannot be grown in the confined volumes of greenhouses and growth chambers. We recently identified an extremely short (20 cm tall) rice line, which is an ideal model for larger rice cultivars. We called this line ‘Super Dwarf’ rice. Here we report the response of Super Dwarf to temperature, photoperiod, photosynthetic photon flux (PPF), and factors that can affect time to head emergence. Vegetative biomass increased 6% per degree Celsius, with increasing temperature from 27 to 31°C. Seed yield decreased by 2% per degree Celsius rise in temperature, and as a result, harvest index decreased from 60 to 54%. The time to heading increased by 2 d for every hour above a 12‐h photoperiod. Yield increased with increasing PPF up to the highest level tested at 1800 μmol m−2 s−1 (12‐h photoperiod; 77.8 mol m−2 d−1). Yield efficiency (grams per mole of photons) increased to 900 μmol m−2 s−1 and then slightly decreased at 1800 μmol m−2 s−1. Heading was delayed by addition of gibberellic acid 3 (GA3) to the root zone but was hastened under mild N stress. Overall, short stature, high yield, high harvest index, and no extraordinary environmental requirements make Super Dwarf rice an excellent model plant for yield studies in controlled environments.

Plant growth during the greenhouse II experiment on the Mir orbital station

We carried out three experiments with Super Dwarf wheat in the Bulgarian/Russian growth chamber Svet (0.1 m 2 growing area) on the Space Station Mir. This paper mostly describes the first of these NASA-supported trials, began on Aug. 13, 1995. Plants were sampled five times and harvested on Nov. 9 after 90 days. Equipment failures led to low irradiance (3, then 4 of 6 lamp sets failed), instances of high temperatures (ca. 37 °C), and sometimes excessive substrate moisture. Although plants grew for the 90 d, no wheat heads were produced. Considering the low light levels, plants were surprisingly green, but of course biomass production was low. Plants were highly disoriented (low light, mirror walls?). Fixed and dried samples and the root module were returned on the U.S. Shuttle Atlantis on Nov. 20, 1995. Samples of the substrate, a nutrient-charged zeolite called Balkanine, were taken from the root module, carefully examined for roots, weighed, dried, and reweighed. The Svet control unit and the light bank were shipped to Moscow. An experiment validation test (EVT) of plant growth and experimental procedures, carried out in Moscow, was highly successful. Equipment built in Utah to measure CO 2, H 2O vapor, irradiance, air and leaf (IR) temperature, O 2, pressure, and substrate moisture worked well in the EVT and in space. After this manuscript was first prepared, plants were grown in Mir with a new light bank and controller for 123 d in late 1996 and 39 days in 1996/1997. Plants grew exceptionally well with higher biomass production than in any previous space experiment, but the ca. 280 wheat heads that were produced in 1996 contained no seeds. Ethylene in the cabin atmosphere was responsible.

Induced abnormality in Mir- and earth grown super dwarf wheat

Super-dwarf wheat grown on the Mir space station using the Svet “Greenhouse” exhibited morphological, metabolic and reproductive abnormalities compared with Earth-grown wheat. Of prominent importance were the abnormalities associated with reproductive ontogeny and the total absence of seed formation on Mir. Changes in the apical meristem associated with transition from the vegetative phase to floral initiation and development of the reproductive spike were all typical of ‘Super-Dwarf’ wheat up to the point of anthesis. Observation of ruptured anthers from the Mir-grown plants revealed what appeared to be normally developed pollen. These pollen gains, however, contained only one nuclei, while normal viable pollen is tri-nucleate. A potentially important difference in the flight experiment, compared with ground reference studies, was the presence of a high level of atmospheric ethylene (1,200 ppb). Ground studies conducted by exposing ‘Super-Dwarf’ wheat to ethylene just prior to anthesis resulted in manifestation of the same abnormalities observed in the space flight samples.

Sensitivity of Wheat and Rice to Low Levels of Atmospheric Ethylene

Ethylene (C2H4) gas is produced throughout the life cycle of plants and can accumulate in closed growth chambers to levels 100 times higher than in outside environments. Elevated atmospheric C2H4 can cause a variety of abnormal responses, but the sensitivity to elevated C2H4 is not well characterized. We evaluated the C2H4 sensitivity of wheat (Triticum aestivum L.) and rice (Oryza sativa L.) in five studies. The first three studies compared the effects of continuous C2H4 levels ranging from 0 to 1000 nmol mol−1 (ppb) in a growth chamber throughout the life cycle of the plants. A short‐term 1000 nmol mol−1 treatment was included in which exposure was stopped at anthesis. Yield was reduced by 36% in wheat and 63% in rice at 50 nmol mol−1 and both species were virtually sterile when continuously exposed to 1000 nmol mol−1 However, the yield reductions were much less with exposure that stopped at anthesis, suggesting the detrimental effect of C2H4 on yield was greatest around the time of seed set. Two additional studies evaluated the differential sensitivity of two wheat cultivars (Super Dwarf and USU‐Apogee) to 50 nmol mol−1 C2H4 at three CO2 levels [350, 1200, 5000 μmol mol−1 (ppm)] in a greenhouse. Yield of USU‐Apogee was not significantly reduced by C2H4 but the yield of Super Dwarf was reduced by 60%. Elevated CO2 did not influence the sensitivity to C2H4 A difference in the C2H4 sensitivity of USU‐Apogee between greenhouse and growth chamber trials suggests that C2H4 sensitivity is dependent on the environment. Collectively, the data suggest that relatively low levels of C2H4 could induce anomalous plant responses by accumulation in greenhouses and growth chambers with inadequate ventilation. The data also suggest that C2H4 sensitivity can be reduced by both genetic and environmental manipulations.

Water stress, CO 2 and photoperiod influence hormone levels in wheat

'Super Dwarf' wheat (Triticum aestivum L.) plants have been grown from seed to maturity in the Mir space station where they were periodically exposed, because of microgravity and other constraints, to water deficit, waterlogging, high CO2 levels, and low light intensities. The plants produced many tillers, but none of them produced viable seed. Studies have been initiated to determine why the plants responded in these ways. In the present study, effects of the listed stresses on abscisic acid (ABA), indole-3-acetic acid (IAA) and isopentenyl adenosine ([9R]iP) levels in roots and leaves of plants grown under otherwise near optimal conditions on earth were measured. Hormones were extracted, purified by HPLC, and quantified by noncompetitive indirect ELISA. In response to water deficit, ABA levels increased in roots and leaves, IAA levels decreased in roots and leaves, and [9R]iP levels increased in leaves but decreased in roots. In response to waterlogging, ABA, IAA and [9R]iP levels briefly increased in roots and leaves and then decreased. When portions of the root system were exposed to waterlogging and/or water deficit, ABA levels in leaves increased while [9R]iP and IAA levels decreased. These responses were correlated with the percentage of the root system stressed. At a low photosynthetic photon flux (100 micromoles m-2 s-1), plants grown in continuous light had higher leaf ABA levels than plants grown using an 18 or 21 h photoperiod.

Differential production of superoxide dismutase and catalase isozymes during infection of wheat by a Fusarium proliferatum -like fungal isolate

AOS are produced during interactions between pathogens and incompatible and compatible plant hosts. Wheat cv. Super Dwarf infected with a Fusarium proliferatum -like species showed increased deposition of lignin-like polymers and dimers of ferulic acid in plant cell wall structures denoting phenolic oxidation. Consequently, we examined whether enzymes connected with AOS metabolism were modified during the interaction between wheat and this Fusarium isolate. Neither SOD nor catalase activities of wheat were altered by Fusarium -infection of sheath tissue. In liquid in vitro cultures, Fusarium produced three anionic SODs, and anionic and cationic catalase isozymes. OneFusarium MnSOD was expressed in infected wheat, while one Cu/ZnSOD and a second MnSOD were not detected. The Cu/ZnSOD was not expressed during growth on carbon- or nitrogen-limited cultures. The fungal anionic catalases that were highly expressed in carbon-limited in vitro culture, but not the cationic catalase, were detected in Fusarium -infected wheat tissue. Thus, specific isozymes of both fungal SOD and catalase appear not to be produced in the wheat sheath. The fungal enzyme expression patterns are consistent with the fungus experiencing nutrient limitation and/or oxidative stress during its process of colonization of the wheat sheath.

Comparative floral development of Mir-grown and ethylene-treated, earth-grown Super Dwarf wheat

To study plant growth in microgravity, we grew Super Dwarf wheat (Triticum aestivum L.) in the Svet growth chamber onboard the orbiting Russian space station, Mir, and in identical ground control units at the Institute of BioMedical Problems in Moscow, Russia. Seedling emergence was 56% and 73% in the two root-module compartments on Mir and 75% and 90% on earth. Growth was vigorous (produced ca. 1 kg dry mass), and individual plants produced 5 to 8 tillers on Mir compared with 3 to 5 on earth-grown controls. Upon harvest in space and return to earth, however, all inflorescences of the flight-grown plants were sterile. To ascertain if Super Dwarf wheat responded to the 1.1 to 1.7 micromoles mol-1 atmospheric levels of ethylene measured on the Mir prior to and during flowering, plants on earth were exposed to 0, 1, 3, 10, and 20 micromoles mol-1 of ethylene gas and 1200 micromoles mol-1 CO2 from 7 d after emergence to maturity. As in our Mir wheat, plant height, awn length, and the flag leaf were significantly shorter in the ethylene-exposed plants than in controls; inflorescences also exhibited 100% sterility. Scanning-electron-microscopic (SEM) examination of florets from Mir-grown and ethylene-treated, earth-grown plants showed that development ceased prior to anthesis, and the anthers did not dehisce. Laser scanning confocal microscopic (LSCM) examination of pollen grains from Mir and ethylene-treated plants on earth exhibited zero, one, and occasionally two, but rarely three nuclei; pollen produced in the absence of ethylene was always trinucleate, the normal condition. The scarcity of trinucleate pollen, abrupt cessation of floret development prior to anthesis, and excess tillering in wheat plants on Mir and in ethylene-containing atmospheres on earth build a strong case for the ethylene on Mir as the agent for the induced male sterility and other symptoms, rather than microgravity.

Main characteristics of biological components of developing life support system observed during the experiments aboard orbital complex MIR.

Since 1990, the orbital complex MIR has witnessed several incubator experiments for determination of spaceflight effects on embryogenesis of Japanese quail. First viable chicks who had completed the whole embryological cycle in MIR microgravity hatched out in 1990; it became clear that newborns would not be able to adapt to microgravity unaided. There were 8 successful incubations of chicks in the period from 1990 to 1999. In 1995-1997 the MIR-NASA space science program united Russian and US investigators. As a result, experiments Greenhouse-1 and 2 were performed with an effort to grow super dwarf wheat from seed to seed, and experiment Greenhouse-3 aimed at receiving two successive generations of Brassica rapa. But results of these experiments could not be used for definitive conclusions concerning effects of spaceflight on plant ontogenesis and, therefore, experiments Greenhouse-4 and 5 were staged within the framework of the Russian national space program. The experiments finally yielded wheat seeds. Some of the seeds was left on the space station and, being planted, gave viable seedlings which, in their turn, produced the second crop of space seeds.

Analysis of the Spaceflight Effects on Growth and Development of Super Dwarf Wheat Grown on the Space Station Mir

The hypothesis being tested is that Super Dwarf wheat, Triticum aestivum L., plants in the Svet Greenhouse onboard the Russian Space Station Mir will complete a life cycle in spaceflight, providing that the environmental conditions necessary for adequate growth on Earth are supplied. Twenty six seeds of wheat were planted in each of 2 rows of 2 root compartments for a total of 104 seeds in Svet. Germination rate at 7 d was 56 and 73% on Mir and 75 and 90% in ground-based controls. Plants were grown throughout the whole cycle of ontogenesis (123 d) with samples gathered at different times to validate the morphological and reproductive stages of the plants. Young plants showed vigorous early seedling growth, with large biomass production, including the formation of 280 floral spikes. Upon return to Earth, comparative analyses showed that the number of tillers and flowers per spikelet were 63.2% and 40% greater, respectively, in Mir-grown plants than in the controls. By contrast, the stem length (52.4%), spike mass (49.2%) and length (23.1%), awn length (75.7%), number of spikelets per spike (42.8%) and number of seeds per spike (100% sterile) from Mir-grown plants were substantially less than the controls. Distribution of moisture and roots throughout the substrate was very good. All florets on Mir-grown spikes ceased development at the same stage of ontogeny. Lack of caryopses formation was attributed to male sterility occurring at different stages of staminal development. Anthers failed to dehisce and pollen grains were smaller and shriveled compared to the controls, suggesting a chronic stress had occurred in the Svet growth chamber. Recent ground-based studies indicated that ethylene, which was measured at 0.3 to 1.8 mg kg-1 in the Mir, almost certainly could have induced male sterility in the wheat plants grown on the Mir.

First flight of the ASTROCULTURE (TM) experiment as a part of the U.S. Shuttle/MIR program.

A number of space-based experiments have been conducted to assess the impact of microgravity on plant growth and development. In general, these experiments did not identify any profound impact of microgravity on plant growth and development, though investigations to study seed development have indicated difficulty in plants completing their reproductive cycle. However, it was not clear whether the lack of seed production was due to gravity effects or some other environmental condition prevailing in the unit used for conducting the experiment. The ASTROCULTURE (TM) flight unit contains a totally enclosed plant chamber in which all the critically important environmental conditions are controlled. Normal wheat (Triticum aestivum L.) growth and development in the ASTROCULTURE (TM) flight unit was observed during a ground experiment conducted prior to the space experiment. Subsequent to the ground experiment, the flight unit was transported to MIR by STS-89, as part of the U.S. Shuttle/MIR program, in an attempt to determine if super dwarf wheat plants that were germinated in microgravity would grow normally and produce seeds. The experiment was initiated on-orbit after the flight unit was transferred from the Space Shuttle to MIR. The ASTROCULTURE (TM) flight unit performed nominally for the first 24 hours after the flight unit was activated, and then the unit stopped functioning abruptly. Since it was not possible to return the unit to nominal operation it was decided to terminate the experiment. On return of the flight unit, it was confirmed that the control computer of the ASTROCULTURE (TM) flight unit sustained a radiation hit that affected the control software embedded in the computer. This experience points out that at high orbital inclinations, such as that of MIR and that projected for the International Space Station, the danger of encountering harmful radiation effects are likely unless the electronic components of the flight hardware are resistant to such impacts.

445 Solid Matrices for Supporting Plant Growth in Space

In preparation for a spaceflight experiment to measure photosynthesis of wheat (PESTO), four solid media were evaluated for use in the rooting modules of the Biomass Production System (BPS), a new plant growth unit for microgravity. The media were commercial peat-vermiculite (PV) mixture, zeolite developed at Johnson Space (Z/JSC), commercial zeolite developed by Boulder Innovative Technologies (Z/BIT), and arcillite (AR) with slow-release fertilizer. Wheat (cv. USU Super Dwarf) was grown in the media at 1500 μmol/mol CO2, 350 μmol·m-2·s-1 PAR, 23 °C, and 75% relative humidity for 18 to 21 days. Water was delivered to the media through porous tubes imbedded in the media, and NDS pressures of -0.1 to -0.5 kPa were maintained with either a static or recirculating standpipe. Plant height, leaf area, and fresh mass were determined for each experiment. Results indicated that the AR and Z/BIT media resulted in larger and more uniform plants than Z/JSC or PV at the same NDS pressure. Additional experiments were conducted with AR to evaluate interactions between particle size and NDS pressure. At ≈14 days after planting, there was a loss of NDS prime in AR >2.0 mm when the NDS pressure was less than -0.3kPa. This resulted in drying of the media and poor plant growth. There was excess water in the media, which resulted in reduced plant size, in AR <1.0 mm at NDS pressures more than -0.3 kPa.

Photosynthetic capacity and dry mass partitioning in dwarf and semi-dwarf wheat ( Triticum aestivum L.)

Efficient use of space and high yields are critical for long-term food production aboard the International Space Station. The selection of a full dwarf wheat (less than 30 cm tall) with high photosynthetic and yield potential is a necessary prerequisite for growing wheat in the controlled, volume-limited environments available aboard long-term spaceflight missions. This study evaluated the photosynthetic capacity and carbon partitioning of a full-dwarf wheat cultivar, Super Dwarf, which is routinely used in spaceflight studies aboard U.S. space shuttle and NASA/Mir missions and made comparisons with other dwarf and semi-dwarf wheat cultivars utilized in other ground-based studies in plant space biology. Photosynthetic capacity of the flag leaf in two dwarf (Super Dwarf, BB-19), and three semi-dwarf (Veery-10, Yecora Rojo, IBWSN 199) wheat cultivars (Triticum aestivum L.) was assessed by measuring: net maximum photosynthetic rate, RuBP carboxylation efficiency, chlorophyll concentration and flag leaf area. Dry mass partitioning of carbohydrates to the leaves, sheaths, stems and ear was also assessed. Plants were grown under controlled environmental conditions in three replicate studies: slightly enriched CO2 (370 micromoles mol-1), high photosynthetic photon flux (1000 micromoles m-2 s-1; 58 mol m-2 d-1) for a 16 h photoperiod, 22/15 degrees C day/night temperatures, ample nutrients and water provided by one-half strength Hoagland's nutrient solution (Hoagland and Arnon, 1950). Photosynthetic capacity of the flag leaf was determined at anthesis using net CO2 exchange rate versus internal CO2 concentration curves measured under saturating light (2000 micromoles m-2 s-1) and CO2 (1000 micromoles mol-1). Dwarf wheat cultivars had greater photosynthetic capacities than the taller semi-dwarfs, they averaged 20% higher maximum net photosynthetic rates compared to the taller semi-dwarfs, but these higher rates occurred only at anthesis, had slightly greater carboxylation efficiencies and significantly increased chlorophyll concentrations per unit leaf area. The reduced-height wheat had significantly less dry mass fraction in the stem but greater dry mass partitioned to the ear than the taller semi-dwarfs (Yecora rojo, IBWSN-199). Studies with detached heads confirm that the head is a significant sink in the shorter wheat cultivars.

Growth and Photosynthetic Responses of Wheat Plants Grown in Space

Growth and photosynthesis of wheat (Triticum aestivum L. cv Super Dwarf) plants grown onboard the space shuttle Discovery for 10 d were examined. Compared to ground control plants, the shoot fresh weight of space-grown seedlings decreased by 25%. Postflight measurements of the O2 evolution/photosynthetic photon flux density response curves of leaf samples revealed that the CO2-saturated photosynthetic rate at saturating light intensities in space-grown plants declined 25% relative to the rate in ground control plants. The relative quantum yield of CO2-saturated photosynthetic O2 evolution measured at limiting light intensities was not significantly affected. In space-grown plants, the light compensation point of the leaves increased by 33%, which likely was due to an increase (27%) in leaf dark-respiration rates. Related experiments with thylakoids isolated from space-grown plants showed that the light-saturated photosynthetic electron transport rate from H2O through photosystems II and I was reduced by 28%. These results demonstrate that photosynthetic functions are affected by the microgravity environment.