The dehydration tolerance of photosynthetic tissues varies greatly along land plant's phylogeny. We investigated the photochemical response to severe dehydration and subsequent rehydration in 318 species representing the major terrestrial biomes and photosynthetic lineages: lichens, bryophytes, pteridophytes and spermatophytes. Using a standardized portable dehydration test with the maximum photochemical efficiency of PSII (Fv/Fm) as indicator, data meeting two key criteria were analyzed: (1) optimal physiological conditions before dehydration and (2) achieving relative water contents (RWC) below 25% at the dehydration stage (with approximately half dropping further to 5%-15%). Lichens and mosses displayed the highest overall Fv/Fm recovery after dehydration, whereas lower tolerance was observed in the other plant groups. Four distinct photochemical responses to dehydration emerged: dehydration-sensitive (61.2%), rehydration-sensitive (18.6%), dehydration-tolerant (13.6%), and dehydration-resistant (6.5%). Pteridophytes and spermatophytes predominantly exhibited dehydration- or rehydration-sensitive responses, whereas dehydration-tolerant dynamics were mainly observed in mosses and resurrection plants. Dehydration-resistant patterns were rare and occurred particularly in polar and high-altitude angiosperms such as Colobanthus quitensis, Deschampsia antarctica and Pozoa coriacea. These findings highlight the evolutionary divergence of dehydration tolerance strategies across the land plant phylogeny and provide insights into the underlying photochemical mechanisms.

Interfacing bioelectronic devices with plants can enable nature-based sensing networks by transducing electrical signals from living plants into real-time environmental data. A key challenge in developing such platforms is to create stable, biocompatible electrodes that provide sufficient adhesion to plant tissue with minimal impedance drift. In this report, an adhesive gel electrode featuring an inkjet-printed poly(3,4-ethylenedioxythiophene) polystyrene sulfonate (PEDOT:PSS) trace on a poly(vinyl alcohol) (PVA) hydrogel substrate with a methyl cellulose adhesive is presented. When attached to the inside of a Venus flytrap (Dionaea muscipula) lobe these biocompatible printed conformable electrodes demonstrate long-term mechanical stability and reliable continuous recording of action potentials, including when removed and reattached over 14 days. Compared to rigid Ag/AgCl electrodes, these bioelectrodes provide higher sensitivity, faster signal dynamics, and comparable signal-to-noise ratios. The gel electrodes maintain low impedance over extended periods (120 hrs), which is critical for long-term monitoring of bioelectronic signals. This stability enabled real-time monitoring of a Venus flytrap's response to environmental factors such as temperature variations, insect activity, and diurnal variations. Additionally, the bioelectrodes were integrated with low-cost ESP32 microcontroller-based electronics, enabling wireless plant-to-plant communication over long distances. This system demonstrates a biologically integrated platform for developing nature-integrated networks that utilize living plants as sensor nodes.

Biogenic volatile organic compounds (BVOCs), primarily emitted into the atmosphere by terrestrial vegetation through biochemical processes, have key ecological functions in protecting vegetation from biotic or abiotic stresses. However, accurately quantifying and predicting changes in BVOC emissions in response to long-term environmental changes large spatial scales remain challenging. The appropriate tools for observing the BVOC emissions at large scales are still missing. Remote sensing of optical signals is a promising solution to fill spatial knowledge gap. We hypothesize that the carotenoid-related vegetation index, such photochemical reflectance index (PRI), is a promising method to investigate BVOCs emitted by plants based on their functional links with carotenoids and photosynthetic activity. We conducted a leaf-level experiment in greenhouse during the summer of 2022 to investigate how the relationships between PRI and BVOC emissions change in response to drought or heat stresses in Scots pine and English oak saplings during the peak of growing season. We aim to address the following questions: (1) What factors control the relationships between PRI and BVOC emissions in response to mild/extreme drought or heat; (2) Will these controlling factors differ between vegetation species or BVOC emission types (e.g., isoprene and monoterpenes)? (3) Can PRI or other carotenoid-related vegetation indices capture the changes of BVOC emissions in response to drought or heat stresses? We will present our preliminary results. The expected outcomes will give new insight into leaf-level mechanistic links between PRI and BVOC emissions for plants in response to climate drought or warming.

Development of a productive, nutritious, and low-input food source is needed for humanity's expanding population on Earth and for long-duration crewed space missions. The exceptionally protein- and micronutrient-rich members of the aquatic plant family Lemnaceae are uniquely suited for both purposes. We here introduce a protocol to accumulate large quantities of the carotenoid zeaxanthin that are retained post-high-light exposure. The function of the essential human micronutrients zeaxanthin, lutein, vitamin E (α-tocopherol), and pro-vitamin A (β-carotene) are briefly reviewed. Attention is given to the role of zeaxanthin in opposing oxidants (reactive oxygen species), the production of which is further enhanced by space radiation. We report a new preharvest finishing procedure – sudden transfer of Lemna plants grown in low light (200 µmol photons m-2 s-1) to high light (1000 µmol photons m-2 s-1) for six hours, which rapidly increased and stabilized zeaxanthin content. Unlike terrestrial crops, Lemna possessed a uniquely high protein content when grown in low light and did not experience any adverse effects of the abrupt five-fold increase in light intensity. These findings provide additional justification for Lemnaceae as food crops with applications on Earth as well as for future long-duration human space missions.

Plants have to cope with different biotic and abiotic stresses, such as the attacks of pathogens and herbivores, high irradiance, droughts, salt stress or nutrient deficiencies throughout their lifetime. These environmental perturbations lead to the regulation of the “primary” and “secondary” carotenoid network. The formers are produced and stored in plastids and they are necessary for photosynthesis and related functions. Otherwise, secondary carotenoids are derivatives of primary carotenoids. They perform other important functions not related to the photosynthetic process but essential for plant survival as regulators of growth and development or as signal molecules at all levels of plant organization. This review provides a complete revision of the status of all these carotenoids, highlighting their newly discovered functions involved in plant-environment interaction. Concurrently, this review covers recent information on how carotenoids perform critical functions for the survival of animals (including humans) and the way they are suitable diagnostic tools for assessing the functioning of terrestrial ecosystems.

Ageing in dry chlorophyllous propagules is leaded by photooxidation through the photosynthetic machinery, but why species differ in longevity and the ageing mechanisms of when light and oxygen are absent are unknown. We hypothesize that the cellular antioxidant capacity is key for the inter- and intra-specific differences in the ageing process. We have tested this hypothesis in chlorophyllous spores of two ferns. They were subjected to four different storage regimes resulting from light/dark and normoxia/hypoxia combinations. Lipophilic and hydrophilic antioxidants, reactive oxygen species (ROS), and photosynthetic pigments were analysed in parallel to germination and the recovery of Fv/Fm over a storage period of up to 22-months. We show that light and oxygen accelerate the ageing process, but their mechanisms (ROS, increase, antioxidant capacity decrease, loss of efficiency of the photosystem II, pigment degradation) appear the same under all conditions tested. The end of the asymptomatic phase of longevity, when a sudden drop of germination occurs, seems to be determined by a threshold in the depletion of antioxidants. Our results support the hypothesis that ageing kinetics in dry plant propagules is determined by the antioxidant system, but also suggests an active role of the photosynthetic machinery during ageing, even in darkness and hypoxia.

The ability of plants to respond to environmental fluctuations is supported by acclimatory adjustments in plant form and function that may require several days and development of a new leaf. We review adjustments in photosynthetic, photoprotective, and foliar vascular capacity in response to variation in light and temperature in terrestrial plants. The requirement for extensive acclimation to these environmental conditions in terrestrial plants is contrasted with an apparent lesser need for acclimation to different light environments, including rapid light fluctuations, in floating aquatic plants for the duckweed Lemna minor. Relevant features of L. minor include unusually high growth rates and photosynthetic capacities coupled with the ability to produce high levels of photoprotective xanthophylls across a wide range of growth light environments without compromising photosynthetic efficiency. These features also allow L. minor to maximize productivity and avoid problems during an abrupt experimental transfer of low-light-grown plants to high light. The contrasting responses of land plants and floating aquatic plants to the light environment further emphasize the need of land plants to, e.g., experience light fluctuations in their growth environment before they induce acclimatory adjustments that allow them to take full advantage of natural settings with such fluctuations.

Atmospheric carbon dioxide (CO2) is projected to be twice as high as the pre-industrial level by 2050. This review briefly highlights key responses of terrestrial plants to elevated CO2 and compares these with the responses of aquatic floating plants of the family Lemnaceae (duckweeds). Duckweeds are efficient at removing CO2 from the atmosphere, which we discuss in the context of their exceptionally high growth rates and capacity for starch storage in green tissue. In contrast to cultivation of terrestrial crops, duckweeds do not contribute to CO2 release from soils. We briefly review how this potential for contributions to stabilizing atmospheric CO2 levels is paired with multiple additional applications and services of duckweeds. These additional roles include wastewater phytoremediation, feedstock for biofuel production, and superior nutritional quality (for humans and livestock), while requiring minimal space and input of light and fertilizer. We, furthermore, elaborate on other environmental factors, such as nutrient availability, light supply, and the presence of a microbiome, that impact the response of duckweed to elevated CO2. Under a combination of elevated CO2 with low nutrient availability and moderate light supply, duckweeds' microbiome helps maintain CO2 sequestration and relative growth rate. When incident light intensity increases (in the presence of elevated CO2), the microbiome minimizes negative feedback on photosynthesis from increased sugar accumulation. In addition, duckweed shows a clear propensity for absorption of ammonium over nitrate, accepting ammonium from their endogenous N2-fixing Rhizobium symbionts, and production of large amounts of vegetative storage protein. Finally, cultivation of duckweed could be further optimized using hydroponic vertical farms where nutrients and water are recirculated, saving both resources, space, and energy to produce high-value products.

Antarctic macroalgal communities of the upper intertidal zone are particularly poor compared with other coastal regions. Exposure to desiccation and freezing combined with the abrasive effect of ice threatens the life of sessile organisms. One of the few species able to colonize this environment is the rhodophyte Pyropia endiviifolia. It belongs to the Bangiales, one of the oldest extant clades of photosynthetic eukaryotes, which has occurred for more than one billion years with basically the same morphological structure. Considering that the extent of Antarctic glaciation is a geologically recent event, we hypothesized that pre-adaptations to desiccation in bangiophytes may have contributed to the success of P. endiviifolia in Antarctica. To test this, we compared its photosynthetic performance and tolerance to desiccation and freezing with those from a related intertidal species, the temperate Atlantic species Porphyra linearis. As evidenced by gas exchange and chlorophyll fluorescence, P. endiviifolia is more adapted to high irradiances than P. linearis. The former species was also more desiccation-tolerant, and showed a higher glass transition temperature. Both species did not differ in chlorophyll content per dry weight, and tolerance to −20°C, even though the ice-nucleation temperature was much higher in P. endiviifolia. Membrane integrity may depend on fatty acid composition in P. endiviifolia, while on enhanced tocopherol in P. linearis. Overall, both species show different strategies to deal with freezing temperatures: supercooling in P. linearis vs. freezing-tolerance in P. endiviifolia. This matches with the probability of being subjected to sub-zero temperatures in their natural environments (lower in the case of P. linearis). Surprisingly, the higher risk of dehydration in the natural habitat of P. linearis is not matched by a higher desiccation tolerance. This observation does not support the initial hypothesis of the study but suggests the opposite: the acclimation to a cold environment results in higher desiccation tolerance.HIGHLIGHTS● Porphyra linearis and Pyropia endiviifolia are remarkably tolerant to desiccation and freezing.● Antarctic P. endiviifolia is remarkably tolerant to desiccation and freezing.● Mechanisms of freezing tolerance could induce a higher tolerance to desiccation.

This review focuses on recently characterized traits of the aquatic floating plant Lemna with an emphasis on its capacity to combine rapid growth with the accumulation of high levels of the essential human micronutrient zeaxanthin due to an unusual pigment composition not seen in other fast-growing plants. In addition, Lemna’s response to elevated CO2 was evaluated in the context of the source–sink balance between plant sugar production and consumption. These and other traits of Lemnaceae are compared with those of other floating aquatic plants as well as terrestrial plants adapted to different environments. It was concluded that the unique features of aquatic plants reflect adaptations to the freshwater environment, including rapid growth, high productivity, and exceptionally strong accumulation of high-quality vegetative storage protein and human antioxidant micronutrients. It was further concluded that the insensitivity of growth rate to environmental conditions and plant source–sink imbalance may allow duckweeds to take advantage of elevated atmospheric CO2 levels via particularly strong stimulation of biomass production and only minor declines in the growth of new tissue. It is proposed that declines in nutritional quality under elevated CO2 (due to regulatory adjustments in photosynthetic metabolism) may be mitigated by plant–microbe interaction, for which duckweeds have a high propensity.

The summer climate in Maritime Antarctica is characterised by high humidity and cloudiness with slightly above zero temperatures. Under such conditions, photosynthetic activity is temperature-limited and plant communities are formed by a few species. These conditions could prevent the operation of the photoprotective xanthophyll (VAZ) cycle as low irradiance reduces the excess of energy and low temperatures limit enzyme activity. The VAZ cycle regulates the dissipation of the excess of absorbed light as heat, which is the main mechanism of photoprotection in plants. To test whether this mechanism operates dynamically in Antarctic plant communities, we characterised pigment dynamics under natural field conditions in two representative species: the moss Polytrichum juniperinum and the grass Deschampsia antarctica. Pigment analyses revealed that the total VAZ pool was in the upper range of the values reported for most plant species, suggesting that they are exposed to a high degree of environmental stress. Despite cloudiness, there was a strong conversion of violaxanthin (V) to zeaxanthin (Z) during daytime. Conversely, the dark-induced enzymatic epoxidation back to V was not limited by nocturnal temperatures. In contrast with plants from other cold ecosystems, we did not find any evidence of overnight retention of Z or sustained reductions in photochemical efficiency. These results are of interest for modelling, remote sensing and upscaling of the responses of Antarctic vegetation to environmental challenges.

Development of a nutritious, sustainable food source is essential to address worldwide deficiencies in human micronutrients. Aquatic floating plants (e.g., species in the family Lemnaceae, duckweeds) are uniquely suited for area-efficient productivity with exceptionally high rates of growth and nutritional quality. Here, we provide an overview of the role of dietary micronutrients (with a focus on carotenoids) in human health and the promise of Lemnaceae as sustainable crops. We examine the effect of growth light environment on plant biomass production and levels of the carotenoids zeaxanthin, lutein, and pro-vitamin A (β-carotene), as well as the antioxidant vitamin E (α-tocopherol), and protein. Data on each of these nutrients are reported on a plant dry biomass basis (as relevant for nutrition) as well as relative to the required input of light energy (as relevant to resource-use efficiency).

Rising atmospheric carbon dioxide (CO2) levels can impact plant photosynthesis and productivity and threaten food security, especially when combined with additional environmental stressors. This study addresses the effects of elevated CO2 in combination with low nutrient supply on Lemna minor (common duckweed). We quantified plant growth rate and nutritional quality (protein content) and evaluated whether any adverse effects of elevated CO2, low nutrients, or the combination of the two could be mitigated by plant-microbe interaction. Plants were grown under controlled conditions and were either uninoculated or inoculated with microorganisms from a local pond that supported L. minor populations. Under low nutrients in combination with high CO2, growth (plant area expansion rate) decreased and biomass accumulation increased, albeit with lower nutritional quality (lower percentage of protein per plant biomass). Inoculation with plant-associated microorganisms restored area expansion rate and further stimulated biomass accumulation while supporting a high protein-to-biomass ratio and, thus, a high nutritional quality. These findings indicate that plant-microbe interaction can support a higher nutritional quality of plant biomass under elevated atmospheric CO2 levels, an important finding for both human and non-human consumers during a time of rapid environmental change.

This study addresses the unique functional features of duckweed via comparison of Lemna gibba grown under controlled conditions of 50 versus 1000 µmol photons m−2 s−1 and of a L. minor population in a local pond with a nearby population of the biennial weed Malva neglecta. Principal component analysis of foliar pigment composition revealed that Malva was similar to fast-growing annuals, while Lemna was similar to slow-growing evergreens. Overall, Lemna exhibited traits reminiscent of those of its close relatives in the family Araceae, with a remarkable ability to acclimate to both deep shade and full sunlight. Specific features contributing to duckweed’s shade tolerance included a foliar pigment composition indicative of large peripheral light-harvesting complexes. Conversely, features contributing to duckweed’s tolerance of high light included the ability to convert a large fraction of the xanthophyll cycle pool to zeaxanthin and dissipate a large fraction of absorbed light non-photochemically. Overall, duckweed exhibited a combination of traits of fast-growing annuals and slow-growing evergreens with foliar pigment features that represented an exaggerated version of that of terrestrial perennials combined with an unusually high growth rate. Duckweed’s ability to thrive under a wide range of light intensities can support success in a dynamic light environment with periodic cycles of rapid expansion.

Summary Desiccation tolerant plants can survive extreme water loss in their vegetative tissues. The fern Anemia caffrorum produces desiccation tolerant (DT) fronds in the dry season and desiccation sensitive (DS) fronds in the wet season, providing a unique opportunity to explore the physiological mechanisms associated with desiccation tolerance. Anemia caffrorum plants with either DT or DS fronds were acclimated in growth chambers. Photosynthesis, frond structure and anatomy, water relations and minimum conductance to water vapour were measured under well‐watered conditions. Photosynthesis, hydraulics, frond pigments, antioxidants and abscisic acid contents were monitored under water deficit. A comparison between DT and DS fronds under well‐watered conditions showed that the former presented higher leaf mass per area, minimum conductance, tissue elasticity and lower CO2 assimilation. Water deficit resulted in a similar induction of abscisic acid in both frond types, but DT fronds maintained higher stomatal conductance and upregulated more prominently lipophilic antioxidants. The seasonal alternation in production of DT and DS fronds in A. caffrorum represents a mechanism by which carbon gain can be maximized during the rainy season, and a greater investment in protective mechanisms occurs during the hot dry season, enabling the exploitation of episodic water availability.

While most ferns avoid freezing as they have a tropical distribution or shed their fronds, wintergreen species in temperate and boreoalpine ecosystems have to deal with sub-zero temperatures. Increasing evidence has revealed overlapping mechanisms of desiccation and freezing tolerance in angiosperms, but the physiological mechanisms behind freezing tolerance in ferns are far from clear. We evaluated photochemical and hydraulic parameters in five wintergreen fern species differing in their ability to tolerate desiccation. We assessed frond freezing tolerance, ice nucleation temperature and propagation pattern, and xylem anatomical traits. Dynamics of photochemical performance and xanthophyll cycle were evaluated during freeze-thaw events under controlled conditions and, in selected species, in the field. Only desiccation-tolerant species, which possessed a greater fraction of narrow tracheids (<18 mu m) than sensitive species, tolerated freezing. Frond freezing occurred in the field at -3.4 +/- 0.9 degrees C (SD) irrespective of freezing tolerance, freezable water content, or tracheid properties. Even in complete darkness, maximal photochemical efficiency of photosystem II was down-regulated concomitantly with zeaxanthin accumulation in response to freezing. This was reversible upon re-warming only in tolerant species. Our results suggest that adaptation for freezing tolerance is associated with desiccation tolerance through complementary xylem properties (which may prevent risk of irreversible cavitation) and effective photoprotection mechanisms. The latter includes de-epoxidation of xanthophylls in darkness, a process evidenced for the first time directly in the field.

This review compares and contrasts the role of carotenoids across the taxa of life-with a focus on the xanthophyll zeaxanthin (and its structural isomer lutein) in plants and humans. Xanthophylls' multiple protective roles are summarized, with attention to the similarities and differences in the roles of zeaxanthin and lutein in plants versus animals, as well as the role ofmeso-zeaxanthin in humans. Detail is provided on the unique control of zeaxanthin function in photosynthesis, that results in its limited availability in leafy vegetables and the human diet. The question of an optimal dietary antioxidant supply is evaluated in the context of the dual roles of both oxidants and antioxidants, in all vital functions of living organisms, and the profound impact of individual and environmental context.

Duckweed is a promising food crop with multiple benefits for space applications. Fresh duckweed could deliver synergistically acting essential antioxidant nutrients to a crew – but only if growth conditions provide the plant with the right cues to trigger antioxidant formation. We grew Lemna gibba under continuous growth light ranging from low to very high intensities (photosynthetic photon flux densities = PPFDs) in order to investigate the effect on plant growth, photosynthesis, and level of carotenoid antioxidants that are essential human micronutrients. Lemna gibba achieved remarkably high growth rates under modest growth PPFD by virtue of superior light absorption resulting from minimal self-shading and high chlorophyll levels. Conversely, L. gibba’s growth rate remained high even under very high growth PPFDs. This notable ability of L. gibba to avoid inactivation of photosynthesis and diminished growth under very high growth PPFDs resulted from a combination of downregulation of chlorophyll synthesis and increased biochemical photoprotection that limited a build-up of excessive excitation energy. This biochemical photoprotection included accumulation of zeaxanthin (an essential human micronutrient) and high levels of zeaxanthin-catalyzed thermal energy dissipation of excess excitation. Compared to the light levels needed to saturate L. gibba photosynthesis and growth, higher light levels were thus required for strong induction of the essential antioxidant zeaxanthin. These results indicate a need for design of light protocols that achieve simultaneous optimization of plant yield, nutritional quality, and light-use efficiency to circumvent the fact that the light requirement to saturate plant growth is lower than that for production of high zeaxanthin levels. How this trade-off between light-use efficiency of growth and nutritional quality might be minimized or circumvented to co-optimize all desired features is discussed.

Summary Resurrection plants recover physiological functions after complete desiccation. Almost all of them are native to tropical warm environments. However, the Gesneriaceae include four genera, remnant of the past palaeotropical flora, which inhabit temperate mountains. One of these species is additionally freezing‐tolerant: Ramonda myconi. We hypothesise that this species has been able to persist in a colder climate thanks to some resurrection‐linked traits. To disentangle the physiological mechanisms underpinning multistress tolerance to desiccation and freezing, we conducted an exhaustive seasonal assessment of photosynthesis (gas exchange, limitations to partitioning, photochemistry and galactolipids) and primary metabolism (through metabolomics) in two natural populations at different elevations. R. myconi displayed low rates of photosynthesis, largely due to mesophyll limitation. However, plants were photosynthetically active throughout the year, excluding a reversible desiccation period. Common responses to desiccation and low temperature involved chloroplast protection: enhanced thermal energy dissipation, higher carotenoid to Chl ratio and de‐epoxidation of the xanthophyll cycle. As specific responses, antioxidants and secondary metabolic routes rose upon desiccation, while putrescine, proline and a variety of sugars rose in winter. The data suggest conserved mechanisms to cope with photo‐oxidation during desiccation and cold events, while additional metabolic mechanisms may have evolved as specific adaptations to cold during recent glaciations.