Plant Signaling & Behavior

Bárány, Ivett; Fadón, Begoña; Risueño, María C.; Testillano, Pilar S.

doi: 10.4161/psb.5.4.11507pmid: 20383055

Plant cell wall polymers are regulated during development, but the specific roles of their different molecular components and the functional meaning of cell wall changes in different cell types and cell processes are still unclear. In the present work the presence and distribution of different cell wall components in Capsicum annuum L. pollen have been analysed in situ in order to monitor how they change during two developmental programmes. These programmes are: pollen development, which is a differentiation process, and stress-induced pollen reprogramming to embryogenesis, which involves proliferation followed later by differentiation processes. Specific antibodies recognizing the major cell wall polymers, the major hemicellulose, xyloglucan (XG), the rhamnogalacturonan II (RGII) pectin domain, and high- and low-methyl-esterified pectins were used for both dot-blot and immunolocalization assays at light and electron microscopy levels during defined developmental stages. For comparison purposes, a similar approach was also used in zygotic embryogenesis and root apical tip growth. Results showed differences in the distribution pattern of these molecular complexes, in the proportion of esterified and de-esterified pectins in the two pollen developmental pathways, and defined wall changes during microspore reprogramming. These changes were associated with proliferation and differentiation events where highly esterified pectins were characteristic of proliferation, while de-esterified pectins, XG and RGII were abundant in walls of differentiating cells. Starch deposits were also studied and the results revealed changes in starch synthesis dynamics after switching the pollen embryogenic developmental programme. These changes occurred together with modifications in the distribution patterns of cell wall polymers, starch accumulation being associated with cell differentiation. As in the case of proliferating cells, esterified pectins were also abundant in the apertures of developing microspores, regions of new cell wall formation., The different distribution patterns of cell wall polymers were common for proliferating cells and differentiating cells in all the plant systems analysed, including zygotic embryos and root tip cells, suggesting that these patterns are markers of proliferation and differentiation events as well as markers of pollen reprogramming to embryogenesis.

Sano, Hiroshi

doi: 10.4161/psb.5.4.10803pmid: 20118668

Since Lamarck proposed the idea of inheritance of acquired traits 200 years ago, much has been said for and against it, but the theory was finally declined after the 1930s. Despite of the negative opinions of the majority of geneticists, botanists and plant breeders have long recognized that altered properties during the growth were occasionally transmitted to the offspring. This was also the case with artificially altered properties such as dwarfism, flowering timing and plant stature, which were induced by a non-mutagenic chemical, 5-azacytidine and its derivatives. As these drugs are powerful inhibitors of DNA methylation in vivo, a close correlation between methylation and phenotypic expression was suggested. Subsequent studies showed that rice plants acquired disease resistance upon demethylation of the corresponding resistant gene, and that both resistant trait and hypomethylated status were inherited by the progeny up to nine generations. Whether or not the methylation pattern changes under natural condition was then questioned, and recent studies have indicated that it indeed naturally changes in response to environmental stresses. Whether or not the altered methylation pattern during the vegetative growth is heritable was also questioned, and studies on toadflax and rice affirmed the question, showing stable maintenance of hypermethylation in the former and hypomethylation in the latter for 250 and 10 years, respectively. The observation strongly suggested that acquired traits can be heritable as far as the acquired methylation pattern is stably transmitted. This concept is consistent with the Lamarck’s theory of the inheritance of acquired traits, which therefore should be carefully reevaluated to reestablish his impaired reputation.

Selosse, Marc Andre; Martos, Florent; Perry, Brian; Maj, Padamsee; Roy, Melanie; Pailler, Thierry

doi: 10.4161/psb.5.4.10791pmid: 20061806

Mycoheterotrophic plants are achlorophyllous plants that obtain carbon from their mycorrhizal fungi. They were usually considered to associate with fungi that are (1) specific of each mycoheterotrophic species and (2) mycorrhizal on surrounding green plants, which are the ultimate carbon source of the entire system. We review here recent works unravelling that some mycoheterotrophic plants are not fungal-specific, and that some mycoheterotrophic orchids associate with saprophytic fungi. A re-examination of ancient data suggests that lower specificity may be less rare than supposed in mycoheterotrophic plants. Association between mycoheterotrophic orchids and saprophytic fungi arose several times in the evolution of the two partners. We speculate that this indirectly illustrates why transition from saprotrophy to mycorrhizal status is common in fungal evolution. Moreover, some unexpected fungi occasionally encountered in plant roots should not be discarded as ‘molecular scraps’, since these facultatively biotrophic encounters may evolve into mycorrhizal symbionts in some other plants.

Kovács-Bogdán, Erika; Nyitrai, Péter; Keresztes, Áron

doi: 10.4161/psb.5.4.10870pmid: 20118667

In contrast to the damaging effect of high-concentration chemical stressors, the same agents in very low (submicromolar) concentrations have a positive effect on the treated plants, which is non-specific (independent of the chemical nature of the agent). The direct responses depend on the treated organ. When leaves are treated, the effects include an increase in chlorophyll content, CO2 fixation, and delaying senescence of chloroplasts. When roots are treated, the direct effect is an increased cytokinin synthesis. This hormone, after being transported to the shoot, exerts secondary effects, which are similar to the primary ones in leaves. The signalization routes involved in the primary effects proved to be the phosphoinositide and MAPK pathways in any stimulated organ. In this mini-review we summarize our current knowledge about the effects of low-concentration stressors and their mechanism of action with the help of the four used model systems: detached non-rooting and rooting leaves, hydroponically treated and sprayed seedlings.

Mandal, Santi M.; Chakraborty, Dipjyoti; Dey, Satyahari

doi: 10.4161/psb.5.4.10871pmid: 20400851

Phenolic acids are the main polyphenols made by plants. These compounds have diverse functions and are immensely important in plant-microbe interactions/ symbiosis. Phenolic compounds act as signaling molecules in the initiation of legume-rhizobia symbioses, establishment of arbuscular mycorrhizal symbioses and can act as agents in plant defense. Flavonoids are a diverse class of polyphenolic compounds that have received considerable attention as signaling molecules involved in plant-microbe interactions compared to the more widely distributed, simple phenolic acids; hydroxybenzoic and hydroxycinnamic acids, which are both derived from the general phenylpropanoid pathway. This review describes the well-known roles attributed to phenolic compounds as nod gene inducers of legume-rhizobia symbioses, their roles in induction of the GmGin1 gene in fungus for establishment of arbuscular mycorrhizal symbiosis, their roles in inducing vir gene expression in Agrobacterium, and their roles as defense molecules operating against soil borne pathogens that could have great implications for rhizospheric microbial ecology. Amongst plant phenolics we have a lack of knowledge concerning the roles of phenolic acids as signaling molecules beyond the relatively well-defined roles of flavonoids. This may be addressed through the use of plant mutants defective in phenolic acids biosynthesis or knock down target genes in future investigations

Abogadallah, Gaber M.

doi: 10.4161/psb.5.4.10873pmid: 20118663

Salt tolerance is a complex trait involving the coordinated action of many gene families that perform a variety of functions such as control of water loss through stomata, ion sequestration, metabolic adjustment, osmotic adjustment and antioxidative defense. In spite of the large number of publications on the role of antioxidative defense under salt stress, the relative importance of this process to overall plant salt tolerance is still a matter of controversy. In this article, the generation and scavenging of reactive oxygen species (ROS) under normal and salt stress conditions in relation to the type of photosynthesis is discussed. The CO2 concentrating mechanism in C4 and CAM plants is expected to contribute to decreasing ROS generation. However, the available data supports this hypothesis in CAM but not in C4 plants. We discuss the specific roles of enzymatic and non enzymatic antioxidants in relation to the oxidative load in the context of whole plant salt tolerance. The possible preventive antioxidative mechanisms are also discussed.

Steffens, Bianka; Sauter, Margret

doi: 10.4161/psb.5.4.10910pmid: 20948297

Waterlogging or flooding are frequently or constitutively encountered by many plant species. The resulting reduction in endogenous O2 concentration poses a severe threat. Numerous adaptations at the anatomical, morphological, and metabolic level help plants to either escape low oxygen conditions or to endure them. Formation of aerenchyma or rapid shoot elongation are escape responses, as is the formation of adventitious roots. The metabolic shift from aerobic respiration to anaerobic fermentation contributes to a basal energy supply at low oxygen conditions. Ethylene plays a central role in hypoxic stress signaling, and G proteins have been recognized as crucial signal transducers in various hypoxic signaling pathways. The programmed death of parenchyma cells that results in hypoxia-induced aerenchyma formation is an ethylene response. In maize, aerenchyma are induced in the absence of ethylene when G proteins are constitutively activated. Similarly, ethylene induced death of epidermal cells that cover adventitious roots at the stem node of rice is strictly dependent on heterotrimeric G protein activity. Knock down of the unique Gα gene RGA1 in rice prevents epidermal cell death. Finally, in Arabidopsis, induction of alcohol dehydrogenase with resulting increased plant survival relies on the balanced activities of a small Rop G protein and its deactivating protein RopGAP4. Identifying the general mechanisms of G protein signaling in hypoxia adaptation of plants is one of the tasks ahead.

Plieth, Christoph

doi: 10.4161/psb.5.4.10717pmid: 20139732

Plants respond to almost any kind of external stimulus with transients in their cytoplasmic free calcium concentration ([Ca2+]c). A huge variety of kinetics recorded by optical techniques has been reported in the past. This variety has been credited the specificity needed to explain how information about incoming stimuli is evaluated by the organism and turned into the right physiological responses which provide advantages for survival and reproduction. A physiological response often takes place away from the site of stimulation. This requires cell-to-cell communication. Hence, responding cells are not necessarily directly stimulated but rather receive an indirect stimulus via cell-to-cell communication. It appears unlikely that the '[Ca2+]c signature' in the primarily stimulated cell is conveyed over long distances via cell-to-cell communication from the 'receptor cells' to the 'effector cells'. Here, a novel aspect is highlighted to explain the variety of [Ca2+] kinetics seen by integrating methods of [Ca2+]c recording. Plants can generally be seen as cellular automata with specific morphology and capable for cell-to-cell communication. Just a few rules are needed to demonstrate how waves of [Ca2+]c-increases percolate through the organism and thereby deliver a broad variety of 'signatures'. Modeling intercellular signaling may be a possible way to find explanations for different kinds of signal transmission, signal amplification, wave formation, oscillations and stimulus-response coupling. The basic examples presented here show that care has to be taken when interpreting cellular '[Ca2+]c signatures' recorded by optical techniques which integrate over a big number of cells or even whole plants.

Paul, Matthew J.; Jhurreea, Deveraj; Zhang, Yuhua; Primavesi, Lucia F.; Delatte, Thierry; Schluepmann, Henriette; Wingler, Astrid

doi: 10.4161/psb.5.4.10792pmid: 20139731

Trehalose 6-phosphate (T6P), the precursor of trehalose, is a signaling molecule in plants with strong effects on metabolism, growth and development. We recently showed that in growing tissues T6P is an inhibitor of SnRK1 of the SNF1-related group of protein kinases1. SnRK1 acts as transcriptional integrator in response to carbon and energy supply. In microarray experiments on seedlings of transgenic Arabidopsis with elevated T6P content we found that expression of SnRK1 marker genes was affected in a manner to be predicted by inhibition of SnRK1 by T6P in vivo1. A large number of genes involved in reactions that utilize carbon, e.g. UDP-glucose dehydrogenase genes involved in cell wall synthesis, were upregulated. T6P was also found to affect developmental signaling pathways, probably in a SnRK1-independent manner. This includes upregulation of genes encoding UDP-glycosyltransferases that are involved in the glycosylation of hormones. In addition, genes involved in auxin response and light signaling were affected. Many of these genes belong to pathways that link the circadian clock to plant growth and development. The overall pattern of changes in gene expression supports a role for T6P in coordinating carbon supply with biosynthetic process involved in growth and development.

Zurbriggen, Matias D.; Carrillo, Néstor; Hajirezaei, Mohammad-Reza

doi: 10.4161/psb.5.4.10793pmid: 20383072

Plants generally react to the attack of non-host and incompatible host microorganisms by inducing pathogenesis-related (PR) genes and localised cell death (LCD) at the site of infection, a process collectively known as the hypersensitive response (HR). Reactive oxygen species (ROS) are generated in various sub-cellular compartments shortly after pathogen recognition, and proposed to cue subsequent orchestration of the HR. Although apoplast-associated ROS production by plasma membrane NADPH oxidases have been most thoroughly studied, recent observations suggest that ROS are generated in chloroplasts earlier in the response and play a key role in execution of LCD. A model is presented in which the initial outcome of successful pathogen detection is ROS accumulation in plastids, likely mediated by mitogen-activated protein kinases and caused by dysfunction of the photosynthetic electron transport chain. ROS signaling is proposed to spread from plastids to the apoplast, through the activation of NADPH oxidases, and from there to adjacent cells, leading to suicidal death in the region of attempted infection.