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Plant biology for space exploration – Building on the past, preparing for the
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DOI: 10.1016/j.lssr.2021.01.003

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 Life Sciences in Space Research 29 (2021) 1–7

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Life Sciences in Space Research

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Review article

Plant biology for space exploration – Building on the past, preparing for
the future
Elizabeth Kordyum a, Karl H. Hasenstein b, *
a
Department of Cell Biology and Anatomy, Institute of Botany NASU, Tereschenkivska Str. 2, 01601 Kiev, Ukraine, United States
b
Biology Department, University of Louisiana at Lafayette, Lafayette, LA, 70504-3602, United States

A R T I C L E I N F O A B S T R A C T

Keywords: A review of past insights of space experiments with plants outlines basic space and gravity effects as well as gene
Space biology expression. Efforts to grow plants in space gradually incorporated basic question on plant productivity, stress
Reproduction response and cultivation. The prospect of extended space missions as well as colonization of the Moon and Mars
Space environment
require better understanding and therefore research efforts on biomass productivity, substrate and water re­
Plant cultivation
lations, atmospheric composition, pressure and temperature and substrate and volume (growth space) re­
quirements. The essential combination of using plants not only for food production but also for regeneration of
waste, and recycling of carbon and oxygen production requires integration of complex biological and engineering
aspects. We combine a historical account of plant space research with considerations for future research on plant
cultivation, selection, and productivity based on space-related environmental conditions.

1. Introduction 2. Plant reproductive biology

Development of Bioregenerative Life Support Systems (BLSS) as
complement of physical and chemical life support systems is necessary
for future space exploration, Martian missions, and the construction of
lunar bases. Higher plants produce edible biomass and oxygen and
recycle water, absorb carbon dioxide, and improve the atmosphere of
closed environments and support psychological health of astronauts.
Because of these multiple functions, plants are essential components of
BLSS (Ferl et al., 2002; Fu et al., 2016; Stankovic, 2018; Wheeler, 2010;
2017). Since the beginning of manned space flight, a wide range of in­
formation on higher plant performance in microgravity has been ob­
tained from experiments performed with angiosperms in vivo and in
vitro (organ, tissue, cell- and protoplast cultures) that were performed
on board of biosatellites, the space shuttle, and orbital stations. The
history of the plant research and design of growth facilities was recently
reviewed (Zabel et al., 2016). Our review focuses on the conceptual
advances derived from previous space and ground experiments and
highlights biological, physical, and practical aspects that need future
investigations.
More than 20 species in seven families of monocots and eudicots
were used in space research; families are classified according to the
APGIII (Haston et al., 2009): Avena sativa L., Hordeum vulgare L., Oryza
sativa L., Triticum aestivum L., T. durum Desf., Zea mays L. (all Poaceae,
monocots); Lens culinaris Medik., Melilotus albus Desf., Pisum sativum L.,
Vigna radiata (L.) R. Wilczek (all Fabaceae, eudicots); Cucumis sativus L.
(Cucurbitaceae, eudicot); Arabidopsis thaliana (L.) Heynh., Brassica rapa
L., Lepidium sativum L. (all Brassicaceae, eudicots); Impatiens balsamina L.
(Balsaminaceae, eudicot); Capsicum annuum L., Solanum tuberosum L.
(both Solanaceae, eudicot); Crepis capillaris (L.) Wallr., Haplopappus
gracilis (Nutt.) A. Gray, Helianthus annuus L., Lactuca sativa L.
Weightlessness does not change the genetically programmed plant
developmental framework; i.e., physiological responses during onto­
genesis show a similar plasticity in space as on Earth. The ability of
plants to grow and develop in orbital flight allows for studying plant
responses to microgravity at different levels of organization and time of
development. Various changes in the structural and functional organi­
zation of cellular organelles, lipid peroxidation and antioxidant activity,
the cell cycle, expression of genes and proteins involved in a wide range
of cellular processes, including Ca2+, lipid and auxin signaling, cell wall
biosynthesis, reactions to stress, general metabolism (primarily

* Corresponding author.
E-mail address: hasenstein@louisiana.edu (K.H. Hasenstein).

https://doi.org/10.1016/j.lssr.2021.01.003
Received 1 December 2020; Received in revised form 5 January 2021; Accepted 16 January 2021
Available online 19 January 2021
2214-5524/© 2021 The Committee on Space Research (COSPAR). Published by Elsevier B.V. This is an open access article under the CC BY-NC-ND license
(http://creativecommons.org/licenses/by-nc-nd/4.0/).
E. Kordyum and K.H. Hasenstein Life Sciences in Space Research 29 (2021) 1–7

carbohydrate and lipid), were described in numerous papers and re­ Table 1
views (Halstead and Dutcher, 1987; Hoson, 2014; Kittang et al., 2014; Experiments with angiosperm plants flowering and fruiting in space flight.
Kordyum et al., 1997; Kordyum and Chapman, 2017; Kwon et al., 2015; Plant Space vehicle/ Main results References
Manzano et al., 2016; Paul et al., 2013a; Vandenbrink and Kiss, 2016; Hardware
Wolverton and Kiss, 2009; Zheng et al., 2015). As a result of studies of Arabidopsis Orbital station Seed-to-seed (Merkys et al.,
the transcriptome and proteome under microgravity, evidence was ob­ thaliana Salyut 7/ Svetoblok development 1984)
tained for the differential expression of genes resulting in organ-specific (L.) Heynh. growth chamber Seed-to-seed (Link et al., 2014,
plant responses (Ferl et al., 2015; Kruse et al., 2020; Paul et al., 2013b, ISS/ADVanced development, second 2003)
AStroCulture (back-to-back) (Yano et al.,
2013c). (ADVASC) growth generation. Smaller 2013)
To understand and prevent negative effects of microgravity, further facility protein bodies in
studies of the development of the generative organs of plants, especially ISS, Kibo–Japanese cotyledons.
fruits and seeds, in space are necessary (Carman et al., 2015; Musgrave, experiment
module/ Plant
2002; Musgrave and Kuang, 2003). As a result of their sessile life and
Experiment Unit
autotrophy, plant reproduction is very diverse. Reproductive biology of (PEU)
plants covers research of mechanisms and processes of sexual and Brassica rapa Orbital station Mir/ Seed-to-seed (Kuang et al.,
asexual reproduction, including transition to reproductive growth, L. cv. Svet growth facility development, second 2000) (Musgrave
flowering, pollination, and fertilization, amphimixis and apomixis, seed Astroplants (back-to-back) et al., 2000) (
generation. Delay of Musgrave and
maturation and dormancy, heterocarpy, seed dispersal and seedling seed development. Kuang, 2001)
recruitment. These stages are being investigated through cell and mo­ Presence of starch
lecular biology, biochemistry, bioinformatics, and statistics. Reproduc­ grains in cotyledons
tive biology also addresses plant sterility, hybridization, and of mature seeds.
Brassica rapa ISS/Biomass Seed-to-seed (Musgrave et al.,
environmental effects such as lack of natural pollinators and ways to
L. cv. Production System development, 2005)
mitigate suboptimal conditions. Astroplants (BPS) immature seeds.
Delay of seed
3. Seed development development.
Asynchrony in
embryo
Plant seed development in microgravity was achieved in different development,
hardware, spaceflights, and experimental conditions (Table 1). Based on changes in embryo
the diversity of experimental evidence, it is safe to state that plants can morphology. Greater
develop successfully in space. quantities of soluble
carbohydrates and
The ability of angiosperm plants to complete vegetative growth and
starch in embryos,
reproduction is very important for space cultivation because specialized suppression of
reproductive organs (such as rhizome, tuber, corm, or bulb) contain protein
reserve starch and proteins – major foodstuffs for astronauts in long- accumulation.
term space flights. Seed quality is “compromised” by development in Brassica rapa Space shuttle Seed-to-seed (Kuang et al.,
L. cv. Columbia, STS-87/ development, 2005, 2000;
microgravity (Kuang et al., 2000; Link et al., 2014; Musgrave et al., Astroplants Plant Growth immature seeds. Popova et al.,
2005; Popova et al., 2002) and apparently mostly by altered Facility Decrease in a size of 2002)
physico-chemical changes as a result of reduced buoyancy and therefore protein bodies in the
insufficient or reduced gas exchange. The main effects of seed devel­ cotyledons. Increase
of starch grains.
opment in space concern 1) delayed seed maturation, 2) altered
Triticum Orbital station Mir/ Formation of sterile (Campbell et al.,
metabolite composition and accumulation rate of seed reserves (protein, aestivum L. Svet growth facility spikes, seed-to-seed 2001; Carman
starch, lipids), embryo morphology and synchrony in embryo develop­ cv. Super development, second et al., 2015;
ment in siliques. Unfortunately, such data are only known for two Dwarf (back-to-back) Levinskikh et al.,
non-edible species of the Brassicaceae family. generation. Decrease 2000, 2001;
in a size and mass of Sychev et al.,
caryopses. 2001; Veselova
4. Vegetative propagation et al., 2003)
cv. Super ISS/Lada growth Seed-to-seed (Baranova et al.,
The possibility of successful vegetative propagation of plants in space Dwarf facility development. Data 2019)
on reserve
has been demonstrated in three spaceflight experiment (Table 2).
substances in grains
It is doubtful that future establishments of plant cultivation facilities are missing.
on Moon or Mars can only rely on generative propagation; rather, the Pisum sativum ISS/Lada growth Four generations. (Sychev et al.,
well-known totipotency of plant cells invites tissue culture based prop­ L. facility Data on reserve 2007)
agation that can more rapidly provide the necessary increase in pro­ substances in grains
are missing.
ductive biomass than seed-derived cultivation techniques. These aspects
are particularly important when seed propagation requires extended
time or special environmental conditions or for woody species that ul­ on fruit and seed development.
timately will have to be part of a plant cultivation system. Performance
and vigor of plants apparently is independent of the propagation method 5. Storage compounds
(Debnath, 2007) but provides opportunities for selection as was
demonstrated for the sterile banana relative Ensete ventricosum (Tesfa­ The nutrient value of plants is mostly determined by their amount of
micael et al., 2020). starch, lipids, vitamins, and minerals. Although some data are available
Plants that have been grown in orbit include two generations each of for starch accumulation in wheat leaves (Stutte et al., 2006), we know
A. thaliana, B. rapa, and T. aestivum and four generations of P. sativum. little about synthesis and accumulation of storage reserves in seeds of
Available data on growth, development, and seed propagation of plants grain legumes and caryopses of cereals under space flight conditions,
in space, indicate that gravity is not required for the development of despite ranking at the top of the list of plant crops recommended for
vegetative part of the plant life cycle, but its absence has adverse effects

2
E. Kordyum and K.H. Hasenstein Life Sciences in Space Research 29 (2021) 1–7

Table 2 performed and led to the development of GeneLab (https://genelab.

Experiments on vegetative propagation of angiosperm plants. nasa.gov/) as a repository for space-related experiments. As of now
Plant Space vehicle/ Main results References genetic analyses are not possible in space and harvested material must
Hardware be returned to earth. This necessity resulted in the development of
Solanum Orbital station Mir/ Formation of tubers. (Kordyum et al., special fixation hardware, the KSC fixation tubes (Paul et al., 2005) and
tuberosum BIOCONTAINER Decrease in a size of 1997; Kordyum, RNA-Later fixation became the standard for space experiments (Ferl
L. cv. growth chamber starch grains. 1997) et al., 2011). However, the often observed up-regulation of diverse stress
Zarevo signals (Choi et al., 2019; Legue et al., 1996; Paul et al., 2001; Paul and
Solanum Space shuttle Formation of tubers. (Brown et al.,
tuberosum Columbia, STS-73/ Increase in the 1997; Cook and
Ferl, 2002) suggests that the RNALater fixation could induce especially
cv. ASTROCULTURE percentage of small Croxdale, 2003; osmotic stress responses in tissue (Park and Hasenstein, 2016).
Norland growth chamber starch grains in Cook et al., Although genetic analyses of space material is common (Correll
tubers. 1998; Croxdale et al., 2013; Ferl et al., 2015; Paul et al., 2012), investigations of
et al., 1997)
organ-specific plant responses to microgravity and cell type-specific
Ipomoea Space shuttle Successful growth of (Mortley et al.,
batatas (L.) Columbia/Plant stem cuttings and 2008) analyses of developmental processes and environmental responses
Lam. cv. growth unit (PGU) regeneration of (Schmidt et al., 2015) are needed. These studies should include the
Whatley/ adventitious roots. development of male and female gametophytes, embryo and endo­
Loretan Substantial increase sperm. Future research on plant reproductive biology will have appli­
in the concentration
of soluble sugars,
cations for general agriculture as well as plant culture in space.
glucose, fructose, Investigations of the fertility of pollen grains and embryo sacs,
sucrose, and total pollination, embryo- and endosperm development, and detection of
starch critical stages in seed development are likely to alleviate reduced per­
formance and yield. Although cell division and differentiation occur
normally in space, deviations in late embryogenesis may lead to devel­
space agriculture (Bugbee and Koemer, 1997; Tibbitts and Alford, 1980;
opmental anomalies. Genomic imprinting or epigenetic modification of
Wheeler, 2017). Despite some physical changes in the starch composi­
parental genomes are not functionally equivalent but well known for the
tion and size of soybean and potato (Kuznetsov et al., 2001), examina­
endosperm of angiosperms (Gehring and Satyaki, 2017; Grossniklaus,
tion of leaf starch content suggested that the spaceflight environment
2005; Kordyum and Mosyakin, 2020; Lopes and Larkins, 1993; Raissig
had minimal impact on wheat starch metabolism.
et al., 2011); however, they have not yet been studied in space. There­
Although lipid content in oil-producing plants has not yet been
fore, the influence of space flight conditions on epigenetic modifications
studied, gene ontology data provide indirect evidence that space flight
of gene expression during the development of the embryo and endo­
affects lipid metabolism (Aubry-Hivet et al., 2014). Interestingly, lipid
sperm should be considered for future experiments. In addition, mo­
degradation during germination in space is delayed (Briarty and Maher,
lecular investigations of generative organs of space-grown plants have
2004) again suggesting that lipid metabolism is affected by space con­
not been conducted. Available information on the effects of space flight
ditions. While direct analyses of vitamin content in plants grown in
only dealt with transcriptomes of vegetative tissue.
space is lacking, clinorotation experiments showed no effect on the
Lastly, it is important to recognize radically different approaches to
vitamin content in leafy vegetables (Rivera et al., 2006), suggesting that
enhance plant performance such as synthetic biology that provides
secondary metabolites are controlled in space by the same factors as on
pathways for adjusting higher plants based on the metabolism and in­
earth.
sights from other organism.(Moses et al., 2017) or specifically adjusting
Because wheat and rice constitute some of the most important food
the expression of specific genes (Xu et al., 2021) to enhance starch
sources on Earth, they most likely will serve that function for future
formation under space conditions.
outposts as well. The regulation of endosperm development, including
possible epigenetic modification as a result of altered environments,
7. Summary
such as lack of gravity, elevated CO2, and other changes in the atmo­
sphere should be studied.
In addition to cereals and legumes, a list of plants recommended for
The synthesis and accumulation of proteins, starch, and lipids is
space agriculture includes a wide range of vegetable crops (Tibbitts and
particularly relevant for balancing storage components in plants. Cur­
Alford, 1980; Wheeler, 2017) that provide a balanced diet and include
rent ground-based research affirms that changes in metabolism between
leaf vegetables and storage roots. Using such crops during long-term
carbohydrate and lipid metabolism have the potential to enhance the
space missions poses the challenges of sustaining their viability.
lipid content of typically starch-producing potatoes (Liu et al., 2017; Xu
Adequate nutrition, maintenance of genetic diversity and reliable seed
et al., 2019). Space-grown plants changed their carbohydrate meta­
stock depends on species-specific studies under space conditions. Un­
bolism in seeds, potato tubers and stem cuttings (Brown et al., 1997;
derstanding plant reproductive biology under space conditions is critical
Kuang et al., 2000; Link et al., 2014; Mortley et al., 2008) but further
for future human settlements on Moon and Mars. Plant development,
characterization of biochemical and cellular mechanisms of storage
adaptation, acclimation and eventually evolution to and in other envi­
accumulation is needed, including specific transport, and packaging, e.
ronments are important research topics for plant scientists. The identi­
g., the formation of protein bodies (Cui et al., 2016; Kumamaru et al.,
fication of mechanisms by which altered gravity, enhanced UV and
2007) in microgravity, especially in the endosperm of cereals and in
ionizing radiation, and other space-related factors affect reproductive
cotyledons of legumes.
structures will be critically important for nutrition and agriculture away
Spaceflight effects on storage reserves are likely the result of stress
from planet Earth. In addition to overcoming space-specific problems,
during cultivation which may decrease quantity, nutritional value, and
such research will improve crop yield on Earth and help mitigate
possibly flavor. However, it is difficult to assess organoleptic properties
climate-change related drought- and temperature stress.
in space because the perception of flavors in space changes (Lane and
Feeback, 2002).
8. Environmental aspects
6. Molecular approaches
Because research is typically focused on a limited set of factors, it is
Molecular analyses of space-grown material are commonly important to consider the complexity of ‘space agriculture’, where
simultaneous occurrences of many extreme and seemingly unrelated

3
E. Kordyum and K.H. Hasenstein
circumstances coincide. Often these factors cannot be simulated on
Earth but are critically important for space applications. The following
compilation highlights some of these issues in hopes that they serve as
guide for future research on space plant biology. The plant-specific
research topics outlined above need to be integrated with space-
specific and non-terrestrial conditions. It is important to distinguish
between the weightlessness environment during space travel and
reduced gravitational conditions on Moon, Mars, or other worlds.
9. Water availability
The first and arguably most important parameter for plant growth
depends on water availability, distribution, and in general the root-
environment interaction. The interaction between water and roots is
dependent on the substrate that surrounds roots, the water content and
root physiology. Because of weight constraints, space applications
would benefit from a ‘soil-less’ culture similar to hydro-, aero- or
aquaponics as it would save mass, a relevant cost factor for launch but a
lesser concern for lunar or Martian bases.
Under weightless conditions, water will flow based on surface in­
teractions and roots likely will be surrounded by layers of water that will
be thick enough to limit gas exchange and lead to hypoxia (Stout et al.,
2001). This condition will have to be controlled either by temporarily
removing excess water or oxygen enrichment as oxygen starvation of
roots can have lethal effects especially during early germination. Star­
vation of O2 can also lead to altered metabolism, especially of malate
(Vitor and Sodek, 2019). Research of space plant cultivation should
include means to control water by non-traditional means. For example,
misting might be useful but the time course of formation of either
droplets or layers too large for root gas exchange in space is unknown. In
terms of substrate, soil rich in organic compounds are desirable but not
available in space, Moon or Mars. Although the pH on Mars was calcu­
lated to be neutral (pH 7.2 ± 0.1) (Plumb et al., 1993), water retention
and capacity are unknown and likely as diverse as on Earth.
10. Reduced pressure
Any agricultural activity needs to take into account the low atmo­
spheric pressure on Mars (0.6 kPa) (Haberle, 2015), and no pressure on
the Moon. The triple point of water (273 K, 611 Pa) implies that either
higher pressure than available on Mars or elevated temperature or a
combination of both are required for liquid water to exist on Mars or at
Mars-like conditions. Such conditions have been studied only for Ara­
bidopsis (Paul et al., 2004; Richards et al., 2006; Zhou et al., 2017),
lettuce and wheat (Corey et al., 1996; Zhang et al., 2015) or bacteria
(Schuerger and Nicholson, 2016; Schuerger et al., 2013) The conditions
chosen to maintain liquid water also affect plant performance, especially
photosynthesis and transpiration. Low temperature reduces shoot
growth but enhances root carbohydrate accumulation (He et al., 2020;
Quadir et al., 2011). The complexity of temperature, pressure, and
reduced gravity interactions is unknown but is likely to affect mineral
and water uptake and metabolite production and distribution. These
parameters should be part of future attempts to optimize plant cultiva­
tion under unusual conditions such as weightlessness or reduced gravity,
elevated CO2, hypobaric conditions, novel substrates without organic
compounds (i.e., regolith), and water scarcity.
11. Atmospheric composition
Plants can communicate via Volatile Organic Compounds (VOCs)
(Matsui, 2016), and especially plant hormones such as ethylene, and
(esters of) salicylic acid and jasmonic acid are known as potent gaseous
regulators of plant growth (Clarke et al., 2000). The realization that the
atmosphere in spacecraft is enriched with numerous VOCs prompted
studies on their effect on plant growth. Perhaps augmented by reduced
buoyancy, tests showed that ethanol concentrations that are deemed
4
Life Sciences in Space Research 29 (2021) 1–7
acceptable for humans (Spacecraft Maximum Allowable Concentration,
SMAC) are problematic for plants as 0.1 SMAC reduces chlorophyll
content, growth and yield and 0.5 SMAC ethanol proved lethal for radish
while acetone did not have negative effects (Stutte et al., 2004).
Although ethylene scrubbers are routinely used for plant space experi­
ments, the variability and sensitivity of plants for VOCs, especially under
space, hypobaric, and reduced temperature conditions are unknown and
require studies.
12. Photoperiodicity
In addition to changes in atmospheric composition, altered photo­
periods need investigation. All evolutionary processes are not just
imprinted by Earth’s gravity profile but also the 24 h daylength, which is
similar to a Mars day (about 24 h and 40 min) but totally different from a
lunar light and dark cycles of about 2754 h each. While extended dark
periods diminish productivity, no information exists as to how long
darkness or extended light periods can be tolerated without negative
effects or whether enhanced intensity or extended photoperiod can
compensate for extended darkness. Most, if not all studies on photo­
periodicity focus on the effect of short- and long-day periods (typically
shorter or longer than 12 h) but studies of extended ratios (e.g., a single
6 h dark period during a three-earth day cycle or 72 h) have not been
done. Such studies would evaluate the tolerance and by inference,
optimization for plant growth and necessary supplemental light for plant
cultivation on the Moon.
The complex interaction of light periods with circadian rhythms and,
in general biological clocks, not only opens new research possibilities
but also provides opportunities to specifically control genes that deter­
mine flowering and therefore generative development (Yang et al.,
2020). The tight connection between CIRCADIAN CLOCK ASSOCI­
ATED1 gene expression and MYB-related transcription factors that are
also affected by phytochrome (Liu et al., 2020) illustrate the vast and
complex interactions between light periodicity, spectral activity and
stress responses (Lai et al., 2012). Periodic responses vary with tem­
perature, humidity and affect phytochemicals (Liebelt et al., 2019) and
the catalytic light activity not only controls elongation growth but also
photomorphogenesis, root architecture, hormonal signaling, and stress
responses. The light-dependent network involves more than 5000 genes
(Bhatnagar et al., 2020) including communication with symbionts and
nodulation (Kong et al., 2020). The complexity of these interactions
documents highly plastic responses that should be studied under space
conditions and especially at different photoperiodicities to determine
the adaptability of plants to extended light periods as well as their
requirement for dark intervals and the effect of modified light spectra.
13. Spectral composition
In addition to the well-known phytochrome system that regulates
flowering (Chory et al., 1996), many other plant-related phenomena
such as shade avoidance (Whitelam and Smith, 1991), circumnutation
(Yoshihara and Iino, 2005), and the balance between photo- and grav­
itropism (Correll and Kiss, 2005), blue and UV light also modify vege­
tative growth and resource distribution between shoot, root, and storage
tissue (Ajdanian et al., 2019; Bukhov et al., 1996; Kong et al., 2018;
Tezuka et al., 1994). Although the higher energy content of blue light
could lead to enhanced photosynthesis, experimental data are incon­
clusive, partially because of higher transpiration as a result of red
light-induced stomatal opening (Wang et al., 2010) for red and blue light
enhanced spectra compared to white light only (Tang et al., 2020).
Previous assessments of effectiveness were based on the continuous
photosynthetically active radiation (PAR) but some variability of red
and far red wavelengths is strong enough to call for a modification of the
PAR to include far-red radiation (Zhen and Bugbee, 2020).
Part of expanded understanding of light control of plant growth in­
cludes effects of UV-A (315 – 400 nm) on the release of volatile
E. Kordyum and K.H. Hasenstein
compounds such as methanol and isoprene (Centritto et al., 2014)
because VOCs in a closed environment such as spacecraft or extrater­
restrial greenhouses, can be problematic. However, the effect of UV is
wavelength dependent such that at least strongly pigmented leaves show
effects on PSII only at shorter wavelengths (367 nm) as reduced
photosynthesis and stomatal conductance, which affects overall growth
(Samuoliene et al., 2020). However, these effects are species and con­
dition specific. UV results in photoinhibition and affects thylakoid
membrane structure and its reduction promotes yield in wheat (Kataria
and Guruprasad, 2015) but UV can enhance stomatal opening (Bocca­
landro et al., 2012), which may also affect water reclamation efforts.
The complexity of assessing light effects comprehensively stems from
(red /far red) light balance to control phytochrome, which promotes
flowering but shortens vegetative growth and increases leaf area and
leaf biomass. In contrast, blue light reduces leaf area but enhances sto­
matal conductance. On the ISS and most likely in any confined and
human-inhabited space environment, the CO2 concentration is elevated
(>3000 ppm) (Georgescu et al., 2020). This elevated concentration not
only affects humans but also plants, especially mesophyll conductance
(Singsaas et al., 2003). Space applications also need to consider the
optimization of canopy height and area; thus, light quality (spectral
composition), fluence (emitted radiation energy) and periodicity
(duration of light and dark cycles) provide many challenges for the
optimization of space agriculture.
14. Ionizing radiation
Human space habitation and travel is strongly affected by ionizing
radiation (IR) but relatively few experiments have tested the effect of IR
on plants. Most of the space radiation consists of energized protons in
the energy range of less than 50 MeV (Durante, 2004). Another type of
ionizing radiation, Galactic Cosmic Rays (GCR) is more energetic but the
prevalence of GCR is inversely proportional to their energy content
(Swordy, 2001). The complex relationship between reduced sensitivity
of seeds in their dry vs. hydrated state, the expansion as a result of water
uptake and thus reduced density of radiation-sensitive nuclei and sec­
ondary effects resulting from the nuclear absorption of high linear en­
ergy transfer (LET) particles, makes predictions of radiation effects on
plants difficult.
After spending five and seven years in space, tomato seeds did not
show noticeable decrease in germination or performance (Marmor and
Martin, 1998) but, interestingly, increased resistance to pathogens
(Mishchenko et al., 2013). Therefore, dry seeds seem fairly resistant to
ionizing radiation in low earth orbit. In contrast, imbibed seeds are more
sensitive (Deoli and Hasenstein, 2018) and can be used for IR-induced
mutagenesis (Tanaka et al., 2010). However, IR can also induce hor­
metic effects that leads to faster germination (Deoli and Hasenstein,
2018; Kim et al., 2005; Okamoto and Tatara, 1995). The lack of
knowledge of the effects of long-term or cumulative exposure to IR on
crop plants requires targeted research on the specific factors such as ion
mass (protons vs. HZE), radiation dose, and cumulative or single dose
exposure. No information is available on cumulative effects of IR on
more than one generation of plants.
15. Conclusion
Research on space effects on plants has two broad emphases: the
plant specific responses and adaptation to altered gravity in space or
Lunar or Martian conditions. Research of plant reproductive biology will
be crucial for the optimization of crop yield under novel conditions. The
second focus concerns investigations of cultivation-limiting effects on
plant growth and performance. We hope that the summary of these two
areas of research will stimulate ground- and space-based work to
improve our understanding of space and extraterrestrial conditions on
plant growth, including breeding efforts to optimize plant performance
and productivity and thus support human requirements and activities.
5
Life Sciences in Space Research 29 (2021) 1–7
Declaration of Competing Interest
None.
Acknowledgement
This work was partially supported through SURA Agreement #
C2019-ULLF-01 to KHH.
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