Production and scavenging of reactive oxygen species and redox signaling during leaf and flower senescence: similar but different

Reactive oxygen species (ROS) play a key role in the regulation of many developmental processes, including senescence, and in plant responses to biotic and abiotic stresses. Several mechanisms of ROS generation and scavenging are similar, but others differ between senescing leaves and petals, despite these organs sharing a common evolutionary origin. Photosynthesis-derived ROS, nutrient remobilization, and reversibility of senescence are necessarily distinct features of the progression of senescence in the two organs. Furthermore, recent studies have revealed specific redox signaling processes that act in concert with phytohormones and transcription factors to regulate senescence-associated genes in leaves and petals. Here, we review some of the recent advances in our understanding of the mechanisms underpinning the production and elimination of ROS in these two organs. We focus on unveiling common and differential aspects of redox signaling in leaf and petal senescence, with the aim of linking physiological, biochemical, and molecular processes. We conclude that the spatiotemporal impact of ROS in senescing tissues differs between leaves and flowers, mainly due to the specific functionalities of these organs

Auxin involvement in tepal senescence and abscission in Lilium: a tale of two lilies

Petal wilting and/or abscission terminates the life of the flower. However, how wilting and abscission are coordinated is not fully understood. There is wide variation in the extent to which petals wilt before abscission, even between cultivars of the same species. For example, tepals of Lilium longiflorum wilt substantially, while those of the closely related Lilium longiflorum×Asiatic hybrid (L.A.) abscise turgid. Furthermore, close comparison of petal death in these two Lilium genotypes shows that there is a dramatic fall in fresh weight/dry weight accompanied by a sharp increase in ion leakage in late senescent L. longiflorum tepals, neither of which occur in Lilium L.A. Despite these differences, a putative abscission zone was identified in both lilies, but while the detachment force was reduced to zero in Lilium L.A., wilting of the fused tepals in L. longiflorum occurred before abscission was complete. Abscission is often negatively regulated by auxin, and the possible role of auxin in regulating tepal abscission relative to wilting was tested in the two lilies. There was a dramatic increase in auxin levels with senescence in L. longiflorum but not in Lilium L.A. Fifty auxin-related genes were expressed in early senescent L. longiflorum tepals including 12 ARF-related genes. In Arabidopsis, several ARF genes are involved in the regulation of abscission. Expression of a homologous transcript to Arabidopsis ARF7/19 was 8-fold higher during senescence in L. longiflorum compared with abscising Lilium L.A., suggesting a conserved role for auxin-regulated abscission in monocotyledonous ethylene-insensitive flowers

Distinctive phytohormonal and metabolic profiles of Arabidopsis thaliana and Eutrema salsugineum under similar soil drying

Main conclusions: Arabidopsis and Eutrema show similar stomatal sensitivity to drying soil. In Arabidopsis, larger metabolic adjustments than in Eutrema occurred, with considerable differences in the phytohormonal responses of the two species. Although plants respond to soil drying via a series of concurrent physiological and molecular events, drought tolerance differs greatly within the plant kingdom. While Eutrema salsugineum (formerly Thellungiella salsuginea) is regarded as more stress tolerant than its close relative Arabidopsis thaliana, their responses to soil water deficit have not previously been directly compared. To ensure a similar rate of soil drying for the two species, daily soil water depletion was controlled to 5–10% of the soil water content. While partial stomatal closure occurred earlier in Arabidopsis (Day 4) than Eutrema (from Day 6 onwards), thereafter both species showed similar stomatal sensitivity to drying soil. However, both targeted and untargeted metabolite analysis revealed greater response to drought in Arabidopsis than Eutrema. Early peaks in foliar phytohormone concentrations and different sugar profiles between species were accompanied by opposing patterns in the bioactive cytokinin profiles. Untargeted analysis showed greater metabolic adjustment in Arabidopsis with more statistically significant changes in both early and severe drought stress. The distinct metabolic responses of each species during early drought, which occurred prior to leaf water status declining, seemed independent of later stomatal closure in response to drought. The two species also showed distinct water usage, with earlier reduction in water consumption in Eutrema (Day 3) than Arabidopsis (Day 6), likely reflecting temporal differences in growth responses. We propose Arabidopsis as a promising model to evaluate the mechanisms responsible for stress-induced growth inhibition under the mild/moderate soil drying that crop plants are typically exposed to

Passeig ecofisiologic per l'espai i pel temps : l'estudi de les alteracions produides pels canvis climatics i atmosferics en l'estructura i el funcionament de les plantes i dels ecosistemes terrestres

En l'estudi de l'impacte del canvi global sobre les plantes i els ecosistemes és necessari estudiar com els gens, les plantes senceres, les espècies i les comunitats interaccionen amb l'ambient, tot duent a terme estudis ecofisiològics a diferents escales espacials i temporals, des de la molècula fins a l'ecosistema, i des de segons fins a segles. Presentem aquí alguns exemples de l'aplicació de l'ecofisiologia a les diferents escales espacials i temporals implicades en l'agenda del canvi global. Explorem alguns resultats recents que revelen la possibilitat d'estimar mitjançant teledetecció els processos fotosintètics a escala de fulla, coberta i ecosistema mitjançant l'índex de reflectància fotoquímic (PRI), un índex basat en un principi en canvis moleculars dels pigments del cicle de les xantofil·les, que pot acabar ajudant a estimar el balanç local, regional i global del carboni en el context del canvi climàtic. Pel que fa a l'aplicació de l'ecofisiologia a diferents escales temporals d'aquest canvi global, presentem alguns exemples d'estudis paleoecofisiològics (milers d'anys), històrics (segles), observacionals (últims anys i dècades), i simuladors (per a les properes dècades), en els quals abordem les interaccions entre les plantes i els diferents components del canvi global canvis en la concentració atmosfèrica de CO2, ozó, òxids de nitrogen, compostos orgànics volàtils, i altres gasos; canvis climàtics; eutrofització de la biosfera, disminució de l'ozó estratosfèric, canvis d'usos del sòl i pèrdua de biodiversitat). Els canvis biològics són sovint espectaculars. Exemples en tenim en l'avançada primavera biològica, els desplaçaments de la vegetació a les muntanyes, la progressiva eutrofització dels ecosistemes o l'augment de les emissions de compostos orgànics volàtils de les plantes, entre molts d'altres.In the study of human and global change impacts on plants and ecosystems, it is necessary to understand how genes, whole plants, species and communities interact with the environment. With this aim, current ecophysiological studies are conducted at different spatial and temporal scales, from molecule to ecosystem, and from seconds to centuries. We present here some examples of the application of ecophysiology at the different spatial and temporal scales involved in the Global Change agenda.We describe some recent results revealing the possibility of remote sensing estimation of photosynthetic processes at leaf, canopy and ecosystem levels by means of the Photochemical Reflectance Index (PRI), an index based in the molecular changes of xanthophylls cycle pigments that can help to estimate local, regional, and global carbon budget. Regarding the application of ecophysiology at different temporal scales of the Global Change, we present some examples of paloeoecophysiological (thousands of years), historical (centuries), observational (last years and decades), and experimental and modelling (next decades) studies, where we approach the interactions between plants and the different components of the Global Change (changes in atmospheric concentration of CO2, ozone, nitrogen oxides, volatile organic compounds, and other gases; climatic changes; biosphere eutrophication; stratospheric ozone decrease; land use changes; and biodiversity loss). Biological changes are often spectacular. Some examples are biological spring advancements, vegetation shifts in the mountains, ecosystems progressive eutrophication, or increases in the emissions of plants volatile organic compounds, among others

Influence of stress history on the response of the dioecious plant Urtica dioica L. to abiotic stress

Background: Reaching maturity is thought to affect plant responses to stress, but few studies have examined thus far how stress periods during the juvenile phase can alter the response of plants in a mature stage, particularly in dioecious plants. Aims: This study tested the hypothesis that sex and stress history can alter the response of Urtica dioica plants to abiotic stress. Methods: Stress indicators (water content, chlorophylls, the F v/F m ratio and the extent of lipid peroxidation) were measured in leaves of juvenile and mature plants exposed to a combination of water and nutrient deficit, with an emphasis on evaluating the effects of stress during the juvenile phase on the stress response of mature plants. Results: Stress treatment during the juvenile phase affected plant response to stress during the mature phase. Leaves of reproductive shoots were the most sensitive to stress history, as shown by increases in lipid peroxidation in leaves of reproductive shoots relative to non-reproductive ones. Leaves of both shoot types responded similarly in males and females, which showed no significant differences in any of the parameters measured. Conclusions: Stress history appears to determine the response of mature U. dioica plants to abiotic stress, plants with a stress history showing acclimation to subsequent stress, leaves of non-reproductive shoots of both males and females being more stress tolerant and allowing plant survival under severe stress

Age and sex-related changes in cytokinins, auxins and abscisic acid in a centenarian relict herbaceous perennial

It is still an unsolved question of fundamental biology if, and how, perennial plants senesce. Here, age- and sex-related changes in phytohormones were tested in Borderea pyrenaica, a small dioecious geophyte relict of the Tertiary with one of the longest lifespan ever recorded for any non-clonal herb (more than 300 years). Biomass allocation, together with levels of cytokinins, auxins and absicisic acid, and other indicators of leaf physiology (chlorophylls, lipid peroxidation and F (v)/F (m) ratio) were measured in juvenile and mature plants, including both males and females of three age classes (up to 50 years, 50-100 years, and over 100 years). Plants maintained intact capacity of their vegetative growth and reproductive potential. Cytokinin levels decreased with age, but only in females. Such sex-related differences, however, were not associated with symptoms of physiological deterioration in leaves, but with an increased reproductive effort in females. It is concluded that B. pyrenaica does not show clear signs of senescence at the organism level. Altered cytokinin levels in females were associated with their reproductive effort, rather than to a degenerative process. The alternate use of five meristematic points in the tuber could explain the extraordinary longevity of this species

Photo-oxidative stress markers reveal absence of physiological deterioration with ageing in Borderea pyrenaica, an extraordinarily long-lived herb

In animals, age-associated disorders are believed to be connected to shifts in the antioxidant/pro-oxidant balance in favour of oxidative stress. However, the contribution of oxidative stress to ageing in long-lived perennials has not been explored to date. Here, we tested age- and sex-related changes in several photo-oxidative stress markers in Borderea pyrenaica, a small dioecious geophyte relict of the Tertiary with one of the longest life spans ever recorded for a non-clonal herb (more than 300years). Given that survival increases with age in B.pyrenaica, we hypothesized that oxidative stress does not increase with ageing because the species develops improved anti-oxidant defence. In three field samplings performed during 2008, 2010 and 2011 in the Central Pyrenees (NE Spain), we examined the effects of ageing and sex on photosynthetic pigment levels, PSII integrity (Fv/Fm ratio), lipid peroxidation, and the extent of photo and anti-oxidant protection in chloroplasts. Furthermore, we explored whether age and sex affect plant response to severe natural desiccation. Both male and female plants maintained chlorophyll levels intact, as well as the Fv/Fm ratio and the levels of lipid peroxidation, irrespective of age. This finding suggests the absence of age-associated oxidative stress at the organismal level. Furthermore, photoprotection mechanisms were found to be similarly efficient in the oldest individuals as in juvenile plants, in terms of xanthophyll cycle de-epoxidation and accumulation of low-molecular-weight antioxidants (carotenoids and tocopherols). Indeed, females over 100years of age were the most resistant to severe desiccation, maintaining higher leaf hydration levels, less chlorophyll degradation and better PSII integrity under stress than females below 100years, males below or above 100years, and juveniles. Synthesis. Neither males nor females of the extraordinarily long-lived B.pyrenaica show age-dependent signs of oxidative stress. This observation suggests that age-induced oxidative stress is not a universal feature of ageing in perennial plants. Indeed, females older than 100years showed signs of negative senescence, in that they registered improved physiological performance with increasing age

No signs of meristem senescence in old Scots pine

Ageing and senescence in plants remain poorly understood. Although meristem totipotency may allow woody perennials to be immortal, relative growth and photosynthetic rates typically decline with age. Trees of ages between 129 and 534years were selected in one of the oldest extant populations of Scots pine. Apical branches were propagated by grafting onto homogeneous juvenile rootstock to eliminate the effects of size and environmental variability and isolate those due to age. The hormonal profile of leaves and seeds along with markers of the physiological status of leaves and their pattern of DNA cytosine methylation were measured 15years after grafting. The percentage of total methylated loci in nuclear DNA increased with increasing meristematic age. However, only very few significant relationships were found between levels of phyto-hormones, pigments or physiological markers either in leaves or seeds and age of the meristem. In addition, shoots grafted from old trees grew as fast as those from younger trees and produced the same number of germinable seeds. Synthesis. We conclude that changes in DNA methylation can occur in old trees. The lack of apparent physiological deterioration in the grafted plants suggests that meristem senescence is not the main factor triggering whole-plant ageing in Scots pine

Changes in phytohormones and oxidative stress markers in buried seeds of Vellozia alata

Little information is still available on the mechanisms underlying seed persistence in the soil in several species, and most particularly in vegetation of the rupestrian fields of the Espinhaco Range in Brazil, where ca. 90% of their species are endemic and are of interest for conservation biology. Here we aimed at examining the putative physiological and biochemical changes seeds of one of this species (Vellozia alata L.B.Sm., Velloziaceae) may experience after burial under natural conditions. Endogenous concentrations of phytohormones and oxidative stress markers were measured in seeds buried in the soil for 12 months. Buried seeds experienced a significant loss of germination capacity, which decreased from 58 to 29% during the first 6 months. This was associated with a decline in gibberellins (by 65%), abscisic acid (by 98%), cytokinins (up to 75%) and jasmonic acid (by 97%) during the first 3 months, while salicylic acid increased at 6 months of burial. Malondialdehyde and tocopherol levels also decreased dramatically to non-detectable values during this period, while all tocotrienol homologues decreased by 30%. By contrast, germination capacity remained constant around 30% between 6 and 12 months of burial. During this period, concentrations of all phytohormones examined remained unaltered, except for salicylic acid, whose levels returned to initial values. Tocotrienols decreased significantly, particularly between 9 and 12 months of burial, while the extent of lipid peroxidation remained constant. It is concluded that in V. alata, (i) seed burial causes dramatic changes in phytohormones and in the extent of lipid peroxidation, while the potential for germination decreases; (ii) after 6 months of burial, seed germination capacity remains constant, at least until one year, which is associated with absence of oxidative damage. (C) 2011 Elsevier GmbH. All rights reserved