Plant Signaling, Behavior and Communication.

Being sessile organisms that need to effectively explore space (above and below ground) and acquire resources through growth, plants must simultaneously consider multiple possibilities and wisely balance the energy they spend on growth with the benefits for survival [...].

The Electrome of a Parasitic Plant in a Putative State of Attention Increases the Energy of Low Band Frequency Waves: A Comparative Study with Neural Systems.

Selective attention is an important cognitive phenomenon that allows organisms to flexibly engage with certain environmental cues or activities while ignoring others, permitting optimal behaviour. It has been proposed that selective attention can be present in many different animal species and, more recently, in plants. The phenomenon of attention in plants would be reflected in its electrophysiological activity, possibly being observable through electrophytographic (EPG) techniques. Former EPG time series obtained from the parasitic plant Cuscuta racemosa in a putative state of attention towards two different potential hosts, the suitable bean (Phaseolus vulgaris) and the unsuitable wheat (Triticum aestivum), were revisited. Here, we investigated the potential existence of different band frequencies (including low, delta, theta, mu, alpha, beta, and gamma waves) using a protocol adapted from neuroscientific research. Average band power (ABP) was used to analyse the energy distribution of each band frequency in the EPG signals, and time dispersion analysis of features (TDAF) was used to explore the variations in the energy of each band. Our findings indicated that most band waves were centred in the lower frequencies. We also observed that C. racemosa invested more energy in these low-frequency waves when suitable hosts were present. However, we also noted peaks of energy investment in all the band frequencies, which may be linked to extremely low oscillatory electrical signals in the entire tissue. Overall, the presence of suitable hosts induced a higher energy power, which supports the hypothesis of attention in plants. We further discuss and compare our results with generic neural systems.

Water Deficit at Vegetative Stage Induces Tolerance to High Temperature during Anthesis in Rice.

BACKGROUND: Crop yields have been affected by many different biotic and abiotic factors. Generally, plants experience more than one stress during their life cycle, and plants can tolerate multiple stresses and develop cross-tolerance. The expected rise in atmospheric CO2 concentration ([CO2]) can contribute to cross-tolerance. Priming is a strategy to increase yield or to maintain yield under stress conditions. Thus, our objective was to evaluate if priming the rice plants with water deficit during the vegetative stage can induce tolerance to heat stress at anthesis and to evaluate the contribution of e[CO2]. METHODS: The experiment was arranged in a completely randomized design in a factorial arrangement. Factor A consisted of the following treatments: water deficit at four-leaf stage (no-stress, and drought stress), heat at anthesis (normal temperature, high temperature), and priming with water deficit at four-leaf stage and heat stress at anthesis; and Factor B was two [CO2] treatments: a[CO2] = 400 ± 40 µmol mol-1 and e[CO2] = 700 ± 40 µmol mol-1. We assessed the effect of the treatments on plant growth, yield, biochemical, and transcriptome alterations. RESULTS: Although e[CO2] affected rice growth parameters, it did not affect the priming effect. Primed plants showed an increase in yield and number of panicles per plant. Primed plants showed upregulation of OsHSP16.9A, OsHSP70.1, and OsHSP70.6. These results showed induced cross-tolerance. CONCLUSIONS: Water deficit at the rice vegetative stage reduces the effect of heat stress at the reproductive stage. Water deficit at the vegetative stage can be used, after further testing in field conditions, to reduce the effect of heat stress during flowering in rice.

Approximate entropy: a promising tool to understand the hidden electrical activity of fruit.

Fruits, like other parts of the plant, appear to have a rich electrical activity that may contain information. Here, we present data showing differences in the electrome complexity of tomato fruits through ripening and discuss possible physiological processes involved. The complexity of the signals, measured through approximate entropy, varied along the fruit ripening process. When analyzing the fruits individually, a decrease in entropy values was observed when they entered the breaker stage, followed by a tendency to increase again when they entered the light red stage. Consequently, the data obtained showed a decrease in signal complexity in the breaker stage, probably due to some physiological process that ends up predominating to the detriment of others. This result may be linked to processes involved in ripening, such as climacteric. Electrophysiological studies in the reproductive stage of the plant are still scarce, and research in this direction is of paramount importance to understand whether the electrical signals observed can transmit information from reproductive structures to other modules of plants. This work opens the possibility of studying the relationship between the electrical activity and fruit ripening through the analysis of approximate entropy. More studies are necessary to understand whether there is a correlation or a cause-response relationship in the phenomena involved. There is a myriad of possibilities for the applicability of this knowledge to different areas, from understanding the cognitive processes of plants to achieving more accurate and sustainable agriculture.

Systemic Signals Induced by Single and Combined Abiotic Stimuli in Common Bean Plants.

To survive in a dynamic environment growing fixed to the ground, plants have developed mechanisms for monitoring and perceiving the environment. When a stimulus is perceived, a series of signals are induced and can propagate away from the stimulated site. Three distinct types of systemic signaling exist, i.e., (i) electrical, (ii) hydraulic, and (iii) chemical, which differ not only in their nature but also in their propagation speed. Naturally, plants suffer influences from two or more stimuli (biotic and/or abiotic). Stimuli combination can promote the activation of new signaling mechanisms that are explicitly activated, as well as the emergence of a new response. This study evaluated the behavior of electrical (electrome) and hydraulic signals after applying simple and combined stimuli in common bean plants. We used simple and mixed stimuli applications to identify biochemical responses and extract information from the electrical and hydraulic patterns. Time series analysis, comparing the conditions before and after the stimuli and the oxidative responses at local and systemic levels, detected changes in electrome and hydraulic signal profiles. Changes in electrome are different between types of stimulation, including their combination, and systemic changes in hydraulic and oxidative dynamics accompany these electrical signals.

High Atmospheric CO2 Concentration Mitigates Drought Effects on Acanthostyles buniifolius an Important Grassland Weed in South America.

The differential growth and yield response of plant species to rising carbon dioxide concentrations and climatic change may alter species diversity within biomes. The Pampa Biome in South America is an important grassland biome of agronomic and environmental importance. Acanthostyles buniifolius (Chirca) is one of the most important weeds in natural pasture areas widely distributed in southern South America and can adversely affect livestock production. The current study was designed to identify possible responses of Chirca to CO2 concentration ([CO2]) and drought that would indicate higher adaptation and potential proliferation within the Pampa Biome. Chirca plants were cultivated at two CO2 concentrations (400 (a[CO2]) and 700 (e[CO2]) µmol mol-1) and two water conditions (under water restriction-15% of the pot capacity; and plants without water restriction-pot capacity). Besides growth parameters, we also determined water potential (ѱw), relative water contents (RWC), proline, glycine betaine, total soluble sugars, hydrogen peroxide, lipid peroxidation, superoxide dismutase (SOD), ascorbate peroxidase (APX) activity, chlorophyll A and B, carotenoids and root dry mass (RDM). Plants exposed to e[CO2] are more efficient in water use and have a greater increase in root dry mass, enabling greater adaptation to climate-induced droughts. Among the biochemical changes observed in the plants under drought stress, the accumulation of proline, glycine betaine, and total soluble sugars were the most evident mechanisms allowing plants to tolerate drought stress by osmotic adjustment.

Do plants pay attention? A possible phenomenological-empirical approach.

Attention is the important ability of flexibly controlling limited cognitive resources. It ensures that organisms engage with the activities and stimuli that are relevant to their survival. Despite the cognitive capabilities of plants and their complex behavioural repertoire, the study of attention in plants has been largely neglected. In this article, we advance the hypothesis that plants are endowed with the ability of attaining attentive states. We depart from a transdisciplinary basis of philosophy, psychology, physics and plant ecophysiology to propose a framework that seeks to explain how plant attention might operate and how it could be studied empirically. In particular, the phenomenological approach seems particularly important to explain plant attention theoretically, and plant electrophysiology seems particularly suited to study it empirically. We propose the use of electrophysiological techniques as a viable way for studying it, and we revisit previous work to support our hypothesis. We conclude this essay with some remarks on future directions for the study of plant attention and its implications to botany.

Stress memory of physiological, biochemical and metabolomic responses in two different rice genotypes under drought stress: The scale matters.

Drought is a pivotal cause for crop yield reductions. When subjected to recurrent external stimuli, plants can develop memory of stress responses that, eventually, enables improved plant tolerance to environmental changes. In addition, despite causal relationships, these responses may vary according to hierarchical levels of observation. Thus, this study aims to check the responses of recurrent and non-recurrent stresses in two rice genotypes observing their drought memory responses at different levels of organization, that is, on a physiological, biochemical and metabolomic scale and for end in global PCA. For this, seventy variables were measured on the scales described in order to obtain a large number of observations. The memory responses were evident in almost all scales observed. The lowland genotype, especially plants not subjected to recurrent water shortage, showed higher damage to the photosynthetic apparatus under drought conditions, although it has exhibited more evident memory response effect after rehydration. On the other hand, the upland genotype appears to be more tolerant to drought insofar lower biochemical damage was observed. Specific behaviors of each genotype at biochemical and metabolomics levels and similar behavior at physiological level were observed. This study demonstrates the importance of observation at different hierarchical levels.

Belowground interactions affect shoot growth in Eucalyptus urophylla under restrictive conditions.

Plant-plant interactions like competition or facilitation between seedlings can have profound implications on their establishment and posterior development. These interactions are variable and depend upon the presence of neighbouring plants and environmental factors. In this work, we studied the effects of the interaction by the roots of Eucalyptus urophylla seedlings from a population under various environmental stressful conditions: water deficit, nutrient deficit, low light, low temperature, and high temperature. To evaluate it, we measured some growth and morphological parameters. We demonstrated that shoot parameters such as the number of leaves, leaf area, and dry weight of the leaves were the most affected parameters due to the belowground plant-plant interaction under various environmental conditions. We did not find evidence for competition among the plants, especially under restrictive conditions. Therefore, the study corroborates the stress-gradient hypothesis, which states that plants' differences under stressful conditions lead to facilitative interactions. It has implications for plant ecology and forestry techniques.

Detection of Different Hosts From a Distance Alters the Behaviour and Bioelectrical Activity of Cuscuta racemosa.

In our study, we investigated some physiological and ecological aspects of the life of Cuscuta racemosa Mart. (Convolvulaceae) plants with the hypothesis that they recognise different hosts at a distance from them, and they change their survival strategy depending on what they detect. We also hypothesised that, as an attempt of prolonging their survival through photosynthesis, the synthesis of chlorophylls (a phenomenon not completely explained in these parasitic plants) would be increased if the plants don't detect a host. We quantified the pigments related to photosynthesis in different treatments and employed techniques such as electrophysiological time series recording, analyses of the complexity of the obtained signals, and machine learning classification to test our hypotheses. The results demonstrate that the absence of a host increases the amounts of chlorophyll a, chlorophyll b, and ß-carotene in these plants, and the content varied depending on the host presented. Besides, the electrical signalling of dodders changes according to the species of host perceived in patterns detectable by machine learning techniques, suggesting that they recognise from a distance different host species. Our results indicate that electrical signalling might underpin important processes such as foraging in plants. Finally, we found evidence for a likely process of attention in the dodders toward the host plants. This is probably to be the first empirical evidence for attention in plants and has important implications on plant cognition studies.

Extended cognition in plants: is it possible?

Plants do not possess brains or neurons. However, they present astonishingly complex behaviors such as information acquisition, memory, learning, decision making, etc., which helps these sessile organisms deal with their ever-changing environments. As a consequence, they have been proposed to be cognitive and intelligent, an idea which is becoming increasingly accepted. However, how plant cognition could operate without a nervous central system remains poorly understood and new insights on this topic are urgently needed. According to the Extended Cognition hypothesis, cognition may also occur beyond the limits of the body, encompassing objects from the environment. This was shown possible in humans and spiders, who actively manipulate their external environment to extend their cognitive capacity. Here, we propose that extended cognition may also be found in plants and could partly explain the complexity of plant behavior. We suggest that plants can extend their cognitive abilities to the environment they manipulate through the root influence zone and the mycorrhizal fungi that associate with them. The possibility of a cognitive process involving organisms from different kingdoms is exciting and worthwhile exploring as it may provide key insights into the origin and evolution of cognition.

Plants as electromic plastic interfaces: A mesological approach.

In this manuscript, we propose that plants are eco-plastic and electromic interfaces that can drive emergent intelligent behaviours from synchronized electrical networks. Behind the semantic and anthropocentric problems related by many authors to the extensive use of the terms cognition, intelligence or even 'consciousness' for plants, we suggest a more pragmatic perspective, considering the vegetal world to be a complex biosystemic entity that is able to co-build the world or a form of the world or of significant reality via a set of reciprocal, emerging and confluent interactions. Speaking of adaptive sensory modalities involving perceptual binding or a global state of receptivity nonlinearly leading to cognitive functions, learning capabilities and intelligent behaviours of plants seem to be the more realistic and operational model to describe how plants perceive and treat environmental data. In this study, we strongly suggest that the electrome, which mainly involves constant spontaneous emission of low voltage potentials, is an early marker and a unifying factor of whole plant reactivity in a constantly changing environment and is therefore the key to understanding the cognitive nature of plants. This dynamic coupling enables plants to be knowledge-accumulating systems that are used by evolution to progress and survive, while mesological plasticity is a unique means for plants to interact as subjects with their milieu (umwelt) or natural habitat and to co-signify a possible world.

Molecular responses to recurrent drought in two contrasting rice genotypes.

MAIN CONCLUSION: The set of variables analyzed as integrated by multivariate analysis of principal components consistently showed a memory effect induced by the drought pre-treatment in AN Cambará plants. The effects of drought can vary ddepending on many factors. Among these the occurrence of a previous water stress may leave a residual effect (memory), influencing the future performance of a plant in response to a new drought event. This study tested the hypothesis that plants experiencing recurrent drought would show more active mechanisms of water deficit tolerance, mainly plants of the genotype that is cultivated often experiencing water shortages periods. Additionally, all the plants subjected to water deficit were rehydrated by 24 h and the expression of transcription factors related to drought responses was re-evaluated. To this end, the water status of two rice genotypes, BRS Querência (flooded) and AN Cambará (dryland), was evaluated to identify molecular alterations likely underpinning drought-memory. In growth stage V5, some plants were exposed to water stress (10% VWC soil moisture-pre-treatment). Thereafter, the pots were rehydrated at the same level as the control pots and maintained under this condition until drought was reapplied (10% VWC) at the reproductive stage (R1-R2). Then, the plants were rehydrated and maintained at pot capacity for 24 h. Overall, the set of variables analyzed integrally by multivariate analysis of principal components consistently showed a memory effect induced by the drought pre-treatment in AN Cambará plants (the dryland genotype). This conclusion, based on data of the biochemical and molecular analyses, was supported by the greater capacity of maintenance of the water status by stomatal regulation of the pre-treated and rehydrated plants after the second drought stimulus.

Tolerance to water deficiency between two soybean cultivars: transgenic versus conventional / Tolerância à deficiência hídrica entre dois cultivares de soja: transgênico versus convencional

The main goal of the study was to evaluate the effects on the development and physiological aspects of soybean plants grown under water deficiency at greenhouse conditions, comparing the levels of tolerance to water deficiency between one transgenic and one conventional cultivar, enabling a systematic way to carried out physiological comparisons between soybean cultivars under drought. The study was divided into completely randomized using a factorial 2×2 design, with five replicates. The experimental design included two replenishment levels of daily irrigation (100 percent and 40 percent), applied to the leaves during the V4 developmental stage (fourth trifoliate fully expanded) and two cultivars, 'CD 202' conventional and 'CD 226RR' transgenic. The results showed that both cultivars had similar effects, caused by water deficiency, on dry mass production, but the transgenic cultivar tended to maintain higher biomass allocation in pods, as well as, higher efficiency of leaves to support dry mass production than conventional cultivar in both water conditions. Moreover, the higher maximum CO2 assimilation values and lower membranes damages in the transgenic cultivar under water deficiency showed that the expected higher drought tolerance of the conventional cultivar was not supported by a more accurate physiological investigation.

O principal objetivo do estudo foi avaliar os efeitos sobre o desenvolvimento e aspectos fisiológicos de plantas de soja cultivadas sob deficiência hídrica em condições de casa de vegetação, comparando os níveis de tolerância à deficiência de água entre um cultivar transgênico e um convencional, gerando uma forma sistemática de realizar comparações fisiológicas entre cultivares de soja sob déficit hídrico. O delineamento experimental foi inteiramente casualizado, em esquema fatorial 2×2, com cinco repetições. Os tratamentos hídricos foram dois níveis de reposição de irrigação diária (100 por cento e 40 por cento), aplicados a partir do estádio V4 (quarto trifólio completamente expandido), em dois cultivares ('CD 202' convencional e 'CD 226RR' transgênico). Os resultados evidenciaram que ambos os cultivares tiveram as mesmas respostas na produção de massa seca, mas o cultivar transgênico tendeu a manter maior alocação de biomassa em vagens, assim como maior eficiência da área foliar para a produção de massa seca em ambas as condições de irrigação. Além disso, os maiores valores de assimilação máxima de CO2 e menores danos às membranas no cultivar transgênico, sob deficiência hídrica, mostraram que a maior tolerância à seca esperada do cultivar convencional não foi apoiada por uma investigação fisiológica mais acurada.

Relação entre adubação fosfatada e deficiência hídrica em soja / Relationship between phosphorus supplying and water deficit in soybean

Este estudo teve como objetivo verificar os efeitos do fósforo sobre a fotossíntese e o crescimento de Glycine max (L.) Merr. da cultivar 'Embrapa 48' sob deficiência hídrica controlada, em condições de casa de vegetação, considerando a hipótese de que suplementações de fósforo poderiam aumentar a tolerância das plantas ao déficit hídrico. Após a formação do primeiro par de folhas totalmente expandidas, foi iniciado o processo de indução de deficiência hídrica, utilizando dois regimes de irrigação, com 100 por cento e 25 por cento de reposição da evapotranspiração. A deficiência hídrica causou reduções significativas no acúmulo de massa seca, no potencial de água foliar, na condutância estomática e na assimilação líquida de CO2 em todos os tratamentos. Os resultados das trocas gasosas indicaram que a suplementação de P, na adubação de G. max da cultivar 'Embrapa 48', resultou em uma redução parcial dos efeitos da deficiência hídrica como suposto inicialmente. Porém, apenas em relação à biomassa do sistema radicular detectou-se algum efeito mitigador do P nas plantas sob deficiência hídrica.

The objective of this study was to investigate the effects of phosphorus supplying on growth and photosynthesis of Glycine max (L.) Merr. cv. 'Embrapa 48', cultivated under water deficit in greenhouse conditions, taking into account the hypothesis that phosphorus suply could improve plant tolerance to drough. After the development of the first pair of totally expanded leaves, it was initiate the process of induction of water deficit, using two irrigation levels, 100 percent and 25 percent of evapotranspiration replacement Water deficit reduced dry mass, leaf water potential, stomatal conduction, and net CO2 assimilation regardless P supply. The results of the photosynthetic analysis indicated that phosphorus supply was just partially effective to reduce water deficit effects on G. max cv. 'Embrapa 48'. However, additional P supply influenced positively the production of plants root biomass under water deficit.

Temporal dynamics of stomatal conductance of plants under water deficit: can homeostasis be improved by more complex dynamics?

Este estudo testou a hipótese de que a condutância estomática de uma população de estômatos em uma folha poderia apresentar um comportamento caótico ou complexo sob diferentes condições hídricas, o que poderia favorecer a capacidade homeostática das plantas. A condutância estomática em folhas de girassol e de beterraba cultivadas com irrigação diária e sob deficiência hídrica foi medida com um analisador de gás por infra-vermelho em condições controladas. Os dados foram registrados a cada 20s durante 6h. As séries temporais obtidas foram analisadas por meio dos coeficientes de Lyapunov, dimensão fractal, entropia KS e complexidade LZ relativa. A condutância estomática nas plantas cultivadas com e sem deficiência hídrica exibiu um comportamento provavelmente caótico. As plantas sob estresse hídrico mostraram uma tendência para um comportamento mais complexo, principalmente as plantas de girassol cuja capacidade homeostática foi superior. Algumas implicações biológicas destes comportamentos são discutidas no texto.