Carmen Arena

Rising daily temperatures and water shortage are two of the major concerns in agriculture. In this work, we analysed the tolerance traits in a tomato line carrying a small region of the Solanum pennellii wild genome (IL12‐4‐SL) when grown under prolonged conditions of single and combined high temperature and water stress. When exposed to stress, IL12‐4‐SL showed higher heat tolerance than the cultivated line M82 at morphological, physiological, and biochemical levels. Moreover, under stress IL12‐4‐SL produced more flowers than M82, also characterized by higher pollen viability. In both lines, water stress negatively affected photosynthesis more than heat alone, whereas the combined stress did not further exacerbate the negative impacts of drought on this trait. Despite an observed decrease in carbon fixation, the quantum yield of PSII linear electron transport in IL12‐4‐SL was not affected by stress, thereby indicating that photochemical processes other than CO2 fixation acted to maintain the electron chain in oxidized state and prevent photodamage. The ability of IL12‐4‐SL to tolerate abiotic stress was also related to the intrinsic ability of this line to accumulate ascorbic acid. The data collected in this study clearly indicate improved tolerance to single and combined abiotic stress for IL12‐4‐SL, making this line a promising one for cultivation in a climate scenario characterized by frequent and long‐lasting heatwaves and low rainfall.

ABSTRACT Climate change is increasing the frequency of high temperature shocks and water shortages, pointing to the need to develop novel tolerant varieties and to understand the mechanisms employed to withstand combined abiotic stresses. Two tomato genotypes, a heat‐tolerant Solanum lycopersicum accession (LA3120) and a novel genotype (E42), previously selected as a stable yielding genotype under high temperatures, were exposed to single and combined water and heat stress. Plant functional traits, pollen viability and physiological (leaf gas exchange and chlorophyll a fluorescence emission measurements) and biochemical (antioxidant content and antioxidant enzyme activity) measurements were carried out. A Reduced Representation Sequencing approach allowed exploration of the genetic variability of both genotypes to identify candidate genes that could regulate stress responses. Both abiotic stresses had a severe impact on plant growth parameters and on the reproductive phase of development. Growth parameters and leaf gas exchange measurements revealed that the two genotypes used different physiological strategies to overcome individual and combined stresses, with E42 having a more efficient capacity to utilize the limiting water resources. Activation of antioxidant defence mechanisms seemed to be critical for both genotypes to counteract combined abiotic stresses. Candidate genes were identified that could explain the different physiological responses to stress observed in E42 compared with LA3120. Results here obtained have shown how new tomato genetic resources can be a valuable source of traits for adaptation to combined abiotic stresses and should be used in breeding programmes to improve stress tolerance in commercial varieties.

Photosynthesis and photoprotective mechanisms were investigated in the field on Laurus nobilis L. and Quercus ilex L. leaves exposed to summer drought (July) and winter cold (February) conditions compared with no-stress conditions (May). In July, net photosynthetic rate (A) and stomatal conductance (gs) decreased significantly compared with May in both species; conversely the highest ETR/A ratio and no difference in

Spent coffee grounds (SCGs), deriving from the high global coffee consumption, can have an appreciable environmental impact. One of the eco-friendly reuses of SCGs is their application as a natural fertilizer. This study adopted a soilless cultivation approach to growing Cucumis sativus seedlings using, as rooting media, different coffee solutions obtained by infusing fresh coffee powder or SCGs from mocha and pods for 7 or 30 days. Seedlings’ germination, eco-physiological and biochemical traits were evaluated and the potential toxicological risks or beneficial effects of SCGs on plants were assessed. Results indicate that, although germination was unaffected, fresh coffee medium inhibited seedling development and that 7 days mocha solution stopped seedling growth and induced leaf structural alterations. Conversely, 30 days mocha and pod suspensions promoted seedling biomass, and increased leaf area, leaf polyphenol concentration, and photochemical efficiency. In conclusion, findin...

In Ranunculus asiaticus L., vernalization of propagation material is a common practice for the production scheduling of cut flowers, however little is known about the plant physiology and metabolism of this species as affected by cold treatments. We investigated the influence of two hybrids, MBO and MDR, and three preparation procedures of tuberous roots, only rehydration (control, C), and rehydration plus vernalization at 3.5 °C for 2 weeks (V2) and for 4 weeks (V4), on plant growth and flowering, leaf photosynthesis, and leaf metabolic profile in plants grown in pot in a cold greenhouse. Net photosynthesis (NP) was higher in MDR than in MBO. In the two genotypes, the NP did not change in V2 and increased in V4 compared to C in MBO, while was unaffected by vernalization in MDR. Quantum yield of PSII electron transport (ΦPSII), linear electron transport rate (ETR) and non-photochemical quenching (NPQ) did not differ in the two hybrids, whereas maximal PSII photochemical efficiency (...

Background and aims Higher plants represent an optimal tool to regenerate resources while producing food in Space. However, the configuration of fertile cultivation substrates based on extraterrestrial resources is still a challenge. We evaluated the adaptability of potato (Solanum tuberosum L., cv. ‘Colomba’) to the growth on six substrates: the MMS-1 Mars regolith simulant, alone (R100) and in mixture with 30% in vol. of green compost (R70C30), a fluvial sand, alone or mixed with 30% of compost (S100 and S70C30), a red soil from Sicily (RS), and a volcanic soil from Campania (VS). Methods We assessed the physicochemical properties of the substrates, and the physiological and biometric parameters of potato plants grown in pot on these substrates, in cold glasshouse. Results Both R100 and S100 were alkaline (pH ≥ 8.6) and coarse-textured, lacking organic matter and pivotal macronutrients. The amendment with compost significantly lowered their alkaline pH and improved the chemical fe...

For deep space exploration, radiation effects on astronauts, and on items fundamental for life support systems, must be kept under a pre-agreed threshold to avoid detrimental outcomes. Therefore, it is fundamental to achieve a deep knowledge on the radiation spatial and temporal variability in the different mission scenarios as well as on the responses of different organisms to space-relevant radiation. In this paper, we first consider the radiation issue for space exploration from a physics point of view by giving an overview of the topics related to the spatial and temporal variability of space radiation, as well as on measurement and simulation of irradiation, then we focus on biological issues converging the attention on plants as one of the fundamental components of Bioregenerative Life Support Systems (BLSS). In fact, plants in BLSS act as regenerators of resources (i.e. oxygen production, carbon dioxide removal, water and wastes recycling) and producers of fresh food. In part...

<p><span>Space exploration always has fascinated scientists, and in the last decades, the interest in this goal has increased exponentially. However, the realization of crewed space exploration missions or the permanence of human settlements on orbital stations or planetary habitats requires Life Support Systems (LSS) based on the interaction among abiotic and biotic elements, resembling terrestrial ecosystems. As on Earth, plants in Space would be involved in air regeneration through photosynthesis, water recovery, waste processing and food production, providing the astronauts with renewed resources and fresh food.</span></p><p><span>The extraterrestrial environment may be considered one the most extreme for organisms’ survival because of several Space factors constraining biological life (e.g., altered gravity, fluid-dynamic and microgravity interaction, modified pressure, temperature). Among them, ionizing radiation influences severely plant growth and metabolic processes.  </span><span>Our study explores the response of tomato (<em>Solanum lycopersicum</em> L. ‘Microtom’), one of most widely cultivated crops, to different doses of sparsely ionizing radiation (IR), with two main goals: 1) to evaluate the morpho-functional mechanisms conferring radioresistance to guarantee their ecological role in closed, controlled environments; 2) to assess the possibility to use IR as a “biostimulant” to produce plant-derived functional food richer in functional compounds.</span></p><p><span>Different doses of X-rays (0.3, 1, 10, 20, and 30 Gy, 6 MV energy), were delivered on dry seeds (DS) and germinated seeds (GS) and compared to not-irradiated controls, to define a dose-response curve and check possible negative/positive outcomes on seedlings. After the irradiation treatments, seeds/seedlings were transferred to a climatic chamber and cultivated under controlled environmental conditions of light, temperature, relative humidity and photoperiod. </span><span>For our study, we adopted a multidisciplinary approach that merges anatomical analyses with measurements of photosynthetic efficiency and biochemical traits, including polyphenols and other antioxidant compounds linked with the nutritional value of derived food. </span><span>The growth and photosynthetic performance of DS and GS plants were followed during the whole plant life cycle, while anatomical traits and fruit antioxidant properties were determined at full maturity of the specific organs. </span></p><p><span>The results showed that the outcomes of radiation are dose-specific and dependent on the irradiated target stage, being SS plants more high-performing in photosynthetic activity and antioxidant content in fruits than SGs. Furthermore, some doses of X-rays act as a booster of bioactive compounds in fruits of both SS and SG plants. </span><span>The outcomes of this research will be helpful to optimize crop production in Space and controlled environment agriculture systems. </span><span>Moreover, the fine analysis of the relations between anatomical, eco-physiological and biochemical traits will furnish valuable insights to understand mechanisms of plant acclimation to stress, useful to manage cultivation factors to improve resource use optimization in controlled environments cultivations on Earth in line with sustainable development goals.</span></p>

Abstract The scenarios for the long-term habitation of space platforms and planetary stations involve plants as fundamental part of Bioregenerative Life Support Systems (BLSS) to support the crew needs. Several constraints may limit plant growth in space: among them ionizing radiation is recognized to severely affect plant cell at morphological, physiological and biochemical level. In this work, plants of Phaseolus vulgaris L. were subjected to four different doses of X-rays (0.3, 10, 50 and 100 Gy) in order to assess the effects of ionizing radiation on this species and to analyze possible mechanisms carried out to overcome the radiation injuries. The effects of X-rays on plant growth were assessed by measuring stem elongation, number of internodes and leaf dry weight. The integrity of photosynthetic apparatus was evaluated by photosynthetic pigment composition and ribulose 1,5-bisphosphate carboxylase (Rubisco) activity, whereas changes in total antioxidant pool and glutathione S transferase activity (GST) were utilized as markers of oxidative stress. The distribution of phenolic compounds in leaf tissues as natural shielding against radiation was also determined. Irradiation of plants at 0.3 and 10 Gy did not determine differences in all considered parameters as compared to control. On the contrary, at 50 and 100 Gy a reduction of plant growth and a decrease in photosynthetic pigment content, as well as an increase in phenolic compounds and a decrease in total antioxidant content and GST activity were found. Only a slight reduction of Rubisco activity in leaves irradiated at 50 and 100 Gy was found. The overall results indicate P. vulgaris as a species with a good potential to face ionizing radiation and suggest its suitability for utilization in BLSSs.

Mediterranean tomato landraces adapted to arid environments represent an option to counteract drought, and to address the complexity of responses to water deficit and recovery, which is a crucial component of plant adaptation mechanisms. We investigated physiological, biochemical and molecular responses of two Mediterranean tomato landraces, ‘Locale di Salina’ (Lc) and ‘Pizzutello di Sciacca’ (Pz) under two dehydration periods and intermediate rehydration in greenhouse pot experiments. Relationship between CO2 assimilation (A) and stomatal conductance under severe water stress (gs < 0.05 mol·m−2·s−1) indicated the occurrence of stomatal and non‐stomatal limitations of photosynthesis. Gas exchange promptly recovered within 2–3 days of rehydration. ABA and gs showed a strict exponential relationship. Both leaf ABA and proline peaked under severe water stress. Lc showed higher accumulation of ABA and higher induction of the expression of both NCED and P5CS genes than Pz. Poly(ADP‐ribose) polymerase increased during imposition of stress, mainly in Lc, and decreased under severe water stress. The two landraces hardly differed in their physiological performance. Under severe water stress, gs showed low sensitivity to ABA, which instead controlled stomatal closure under moderate water stress (gs > 0.15 mol·m−2·s−1). The prompt recovery after rehydration of both landraces confirmed their drought‐tolerant behaviour. Differences between the two landraces were instead observed at biochemical and molecular levels.

The growth of plants in Space is a fundamental issue for Space exploration. Plants play an important role in the Bioregenerative Life Support Systems (BLSS) to sustain human permanence in extraterrestrial environments. Under this perspective, plants are basic elements for oxygen and fresh food production as well as air regeneration and psychological support to the crew. The potentiality of plant survival and reproduction in space is limited by the same factors that act on the earth (e.g. light, temperature and relative humidity) and by additional factors such as altered gravity and ionizing radiation. This paper analyzes plant responses to space radiation which is recognized as a powerful mutagen for photosynthetic organisms thus being responsible for morpho-structural, physiological and genetic alterations. Until now, many studies have evidenced how the response to ionizing radiation is influenced by several factors associated both to plant characteristics (e.g. cultivar, species, developmental stage, tissue structure) and/or radiation features (e.g. dose, quality and exposure time). The photosynthetic machinery is particularly sensitive to ionizing radiation. The severity of the damages induced by ionizing radiation on plant cell and tissues may depend on the capability of plants to adopt protection mechanisms and/or repair strategies. In this paper a selection of results from studies on the effect of ionizing radiations on plants at anatomical and eco-physiological level is reported and some aspects related to radioresistance are explored.

The effects of the irrigation regime and different fertilizer sources on the eco-physiological responses and yield characteristics of dragon’s head were explored in a factorial experiment based on a randomized complete block design with 12 treatments and 3 replications in the 2019 growing season. The treatments included six different fertilizer sources (animal manure, vermicompost, poultry manure, biofertilizer, chemical fertilizer, and control) and two irrigation regimes (rainfed and supplemental irrigation). The results indicated the positive effects of supplementary irrigation and the application of vermicompost, poultry manure, and animal manure by increasing the absorption of nutrients (phosphorus and potassium) and improving relative water contents, chlorophyll and carotenoid contents, and the fixed oil percentage of dragon’s head. The activities of catalase, ascorbate peroxidase, and superoxide dismutase decreased in the rainfed plants, whereas organic fertilizer application ...

An adequate and balanced diet is fundamental in preserving the health of astronauts from several space-induced diseases. Therefore, the integration of a diet with fresh food, rich in bioactive compounds such as microgreens produced directly onboard, may be useful in space for human nutrition. However, ionizing radiation (IR) in space represents a significant hindrance for organisms, with potential critical outcomes on plant morpho-anatomical, eco-physiological, and biochemical aspects, depending on the plant and IR features (e.g., species, developmental stage, IR dose, and type). In this study, we analyzed the effect of different doses of X-rays (0-control, 0.3, 1, 10, 20, and 30 Gy) on the morpho-anatomical and nutritional traits of microgreens of Brassica rapa L., irradiated at the stage of germinated seeds. After the irradiation, microgreens were cultivated in controlled conditions. At harvest, the morpho-biometric traits were analyzed, along with the leaf functional anatomical t...

The consumption of fresh tomatoes and processed tomato products is widespread in the Mediterranean diet. This fruit is a valuable source of antioxidants and plays an important role in preventing oxidative stress. This study aimed to investigate the content of antioxidants and measure the total antioxidant capacity (ABTS and DPPH assays) in the peel, pulp, and seed fractions of six tomato cultivars. Finally, some bioactive compounds and total antioxidant activity were also determined in homemade tomato purees, since such homemade production is commonplace in Southern Italy. The level of antioxidants and total antioxidant capacity in each fraction were also calculated based on their actual fresh weight in the whole tomato. The overall results indicated that the peel and seeds of all analysed tomato cultivars contribute significantly to the antioxidant charge of the fruits. Consequently, consuming tomatoes without peel and seeds results in a substantial loss of compounds beneficial for...

Ionising radiation may have different effects on plant metabolism, growth and reproduction, depending on radiation dose, plant species, developmental stage and physiological traits. In this study, exposure of dwarf bean plants to different doses of X-rays (0.3, 10, 50, 100 Gy) was investigated with a multidisciplinary approach consisting of morphological, ecophysiological and biochemical analysis. Both mature and young leaves still growing during the X-rays exposure were compared with non-irradiated control leaves. In particular, leaf expansion, leaf anatomy and functional traits, as well as photosynthetic pigment content and Rubisco expression were analysed. Moreover, the activity of poly(ADP-ribose) polymerase (PARP) was also measured as an indicator of radiation-induced DNA damage. Our data showed that leaf growth is affected by high levels of radiation and demonstrate that mature leaves are more radio-resistant than young leaves, which experience severe dose-dependent changes in leaf functional traits. In particular, young leaves exhibited a reduction of area and an increase in specific mass and dry matter content, as well as a decline in Rubisco activity. Moreover, they showed elevated PARP activity and an increase in phenolic compounds in wall cells if compared with mature leaves. Both of these strategies have been interpreted as a way to help developing leaves withstand irradiation.

Positive stress or essential and nonessential elements can improve nutritive values (biofortification) of edible plants. In the present study, we evaluate (i) the effect of moderate salinity on lettuce biofortification, evaluated as nutritional bioactive compound accumulation, and (ii) the role of iodine in enhancing salt tolerance by increasing photorespiration and the content of antioxidants in lettuce. Physiological (gas exchange and chlorophyll fluorescence emission) and biochemical (photosynthetic pigment and bioactive compound) analyses were performed on lettuce plants grown under moderate salinity (50 mM NaCl alone or 50 mM NaCl in combination with iodine, KIO3). Our results show that NaCl + iodine treatment improves the nutritional value of lettuce in terms of bioactive compounds acting as antioxidants. More specifically, iodine enhances the accumulation of photosynthetic pigments and polyphenols, such as anthocyanins, under salt but does not improve the salt tolerance. Our ...

Plants are able to acclimate to environmental constraints through functional modifications that may also occur in tissues that are not directly exposed to stress. This process is termed “systemic acquired acclimation.” The present study aims to evaluate the involvement of PolyADP-ribose) polymerase (PARP) protein in the acclimation process to high light (HL) stress in Phaseolus vulgaris plants. For this purpose, some leaves located at the top of the plant, in the apical position, were directly exposed to HL (“inducing” leaves), while others on the same plant, distal from the top, continued to be exposed to growth light (“receiving” leaves) to verify the hypothesis that an “alert” message may be transferred from injured tissues to distal ones. Biochemical and eco-physiological analyses, namely PARP activity, H2O2 and water- and fat-soluble antioxidants (i.e., ascorbic acid, tocopherol, glutathione (GSH), phenols, carotenoids, etc.) content, and chlorophyll fluorescence measurements w...

Beta vulgaris L. is a crop selected for cultivation in Space for its nutritional properties. However, exposure to ionizing radiation (IR) can alter plant photosynthetic performance and phytochemical production in the extraterrestrial environment. This study investigated if plant growth under different light quality regimes (FL—white fluorescent; RGB—red–green–blue; RB—red–blue) modifies the photosynthetic behavior and bioactive compound synthesis of plants sprouted by dry seeds irradiated with carbon or titanium high-energy ions. The study evidenced that: (i) the plant response depends on the type of heavyion; (ii) control and C-ion-irradiated plants were similar for photosynthetic pigment content and PSII photochemical efficiency, regardless of the LQ regime; (iii) under FL, net photosynthesis (AN) and water use efficiency (iWUE) declined in C- and Ti-ion plants compared to control, while the growth of irradiated plants under RGB and RB regimes offset these differences; (iv) the in...

The abuse of artificial food dyes and the evidence that they harm human health recently prompted a significant effort to introduce vegan substitutes prepared from fruits and vegetables. Not much information, however, has been collected on their possible effects on aquatic and terrestrial ecosystems once released as waste in surface waters. For this purpose, we analyzed the effects of a vegan red (VEG) preparation (concentration 1.2 g/L) on three rapidly proliferating models for terrestrial and aquatic ecosystem contamination. In particular, in vitro cells cultures (exposure for 24 h), Artemia salina nauplii and Cucumis sativus seedlings (exposure 5 days). A comparison was made with the effects exerted by the two dyes that vegan red is intended to replace: an animal dye, cochineal E120 and an artificial dye E124. The analyses of conventional endpoints, indicative of cell proliferation, differentiation, and growth rate, demonstrate that the three dyes affect development and that the v...

The growing environmental spreading of food synthetic dyes and bio-colors have the potential for altering organisms’ redox states. Here, three model species for aquatic pollution trials, Cucumis sativus seeds, Artemia salina cysts, and Danio rerio embryos, were short-term exposed to a fixed concentration of the artificial red E124, and two red bio-colors, cochineal E120, and vegan red (VEGR). In the animal models, we evaluated the total reactive oxygen species (ROS) and the susceptibility to in vitro oxidative stress, and in C. sativus, H2O2 production and antioxidant capacity. We also measured organismal performance indices (routine oxygen consumption in the animal models, dark oxygen consumption, and photosynthetic efficiency in C. sativus). In C. sativus, only E124 increased ROS and affected dark oxygen consumption and photosynthetic efficiency, while all dyes enhanced the antioxidant defenses. In the A. salina nauplii, all dyes increased ROS, while E120 and E124 reduced the susc...

Light quality plays an essential role in setting plant structural and functional traits, including antioxidant compounds. This paper aimed to assess how manipulating the light spectrum during growth may regulate the photosynthetic activity and fruit bioactive compound synthesis in Solanum lycopersicum L. cv. ‘Microtom’ to improve plant physiological performance and fruit nutritional value. Plants were cultivated under three light quality regimes: red-green-blue LEDs (RGB), red-blue LEDs (RB) and white fluorescent lamps (FL), from sowing to fruit ripening. Leaf functional traits, photosynthetic efficiency, Rubisco and D1 protein expression, and antioxidant production in fruits were analyzed. Compared to FL, RGB and RB regimes reduced height and increased leaf number and specific leaf area, enhancing plant dwarf growth. The RGB regime improved photosynthesis and stomatal conductance despite lower biomass, favoring Rubisco synthesis and carboxylation rate than RB and FL regimes. The RB...

The increase in severe drought events due to climate change in the areas traditionally suitable for viticulture is enhancing the need to understand how grapevines regulate their photosynthetic metabolism in order to forecast specific cultivar adaptive responses to the changing environment. This study aims at evaluating the association between leaf anatomical traits and eco-physiological adjustments of the ‘Falanghina’ grapevine under different microclimatic conditions at four sites in southern Italy. Sites were characterized by different pedoclimatic conditions but, as much as possible, were similar for plant material and cultivation management. Microscopy analyses on leaves were performed to quantify stomata and vein traits, while eco-physiological analyses were conducted on vines to assess plant physiological adaptation capability. At the two sites with relatively low moisture, photosynthetic rate, stomatal conductance, photosystem electron transfer rate, and quantum yield of PSII...