The auxin influx carrier is essential for correct leaf positioning

Auxin is of vital importance in virtually every aspect of plant growth and development, yet, even after almost a century of intense study, major gaps in our knowledge of its synthesis, distribution, perception, and signal transduction remain. One unique property of auxin is its polar transport, which in many well-documented cases is a critical part of its mode of action. Auxin is actively transported through the action of both influx and efflux carriers. Inhibition of polar transport by the efflux inhibitor N-1-naphthylphthalamic acid (NPA) causes a complete cessation of leaf initiation, a defect that can be reversed by local application of the auxin, indole-3-acetic acid (IAA), to the responsive zone of the shoot apical meristem. In this study, we address the role of the auxin influx carrier in the positioning and outgrowth of leaf primordia at the shoot apical meristem of tomato. By using a combination of transport inhibitors and synthetic auxins, we demonstrate that interference with auxin influx has little effect on organ formation as such, but prevents proper localization of leaf primordia. These results suggest the existence of functional auxin concentration gradients in the shoot apical meristem that are actively set up and maintained by the action of efflux and influx carriers. We propose a model in which efflux carriers control auxin delivery to the shoot apical meristem, whereas influx and efflux carriers regulate auxin distribution within the meristem.

No shift to a deeper water uptake depth in response to summer drought of two lowland and sub-alpine C3-grasslands in Switzerland

Temperate C3-grasslands are of high agricultural and ecological importance in Central Europe. Plant growth and consequently grassland yields depend strongly on water supply during the growing season, which is projected to change in the future. We therefore investigated the effect of summer drought on the water uptake of an intensively managed lowland and an extensively managed sub-alpine grassland in Switzerland. Summer drought was simulated by using transparent shelters. Standing above- and belowground biomass was sampled during three growing seasons. Soil and plant xylem waters were analyzed for oxygen (and hydrogen) stable isotope ratios, and the depths of plant water uptake were estimated by two different approaches: (1) linear interpolation method and (2) Bayesian calibrated mixing model. Relative to the control, aboveground biomass was reduced under drought conditions. In contrast to our expectations, lowland grassland plants subjected to summer drought were more likely (43–68 %) to rely on water in the topsoil (0–10 cm), whereas control plants relied less on the topsoil (4–37 %) and shifted to deeper soil layers (20–35 cm) during the drought period (29–48 %). Sub-alpine grassland plants did not differ significantly in uptake depth between drought and control plots during the drought period. Both approaches yielded similar results and showed that the drought treatment in the two grasslands did not induce a shift to deeper uptake depths, but rather continued or shifted water uptake to even more shallower soil depths. These findings illustrate the importance of shallow soil depths for plant performance under drought conditions.

Aluminum toxicity in a tropical montane forest ecosystem in southern Ecuador

Aluminum phytotoxicity frequently occurs in acid soils (pH < 5.5) and was therefore discussed to affect ecosystem functioning of tropical montane forests. The susceptibility to Al toxicity depends on the sensitivity of the plant species and the Al speciation in soil solution, which can vary highly depending e.g., on pH, ionic strength, and dissolved organic matter. An acidification of the ecosystem and periodic base metal deposition from Saharan dust may control plant available Al concentrations in the soil solutions of tropical montane rainforests in south Ecuador. The overall objective of my study was to assess a potential Al phytotoxicity in the tropical montane forests in south Ecuador. For this purpose, I exposed three native Al non-accumulating tree species (Cedrela odorata L., Heliocarpus americanus L., and Tabebuia chrysantha (Jacq.) G. Nicholson) to increased Al concentrations (0 – 2400 μM Al) in a hydroponic experiment, I established dose-response curves to estimate the sensitivity of the tree species to increased Al concentrations, and I investigated the mechanisms behind the observed effects induced by elevated Al concentrations. Furthermore, the response of Al concentrations and the speciation in soil solution to Ca amendment in the study area were determined. In a final step, I assessed all major Al fluxes, drivers of Al concentrations in ecosystem solutions, and indicators of Al toxicity in the tropical montane rainforest in Ecuador in order to test for indications of Al toxicity. In the hydroponic experiment, a 10 % reduction in aboveground biomass production occurred at 126 to 376 μM Al (EC10 values), probably attributable to decreased Mg concentrations in leaves and reduced potosynthesis. At 300 μM Al, increased root biomass production of T. chrysantha was observed. Phosphorus concentrations in roots of C. odorata and T. chrysantha were significantly highest in the treatment with 300 μM Al and correlated significantly with root biomass, being a likely reason for stimulated root biomass production. The degree of organic complexation of Al in the organic layer leachate, which is central to plant nutrition because of the high root density, and soil solution from the study area was very high (mean > 99 %). The resulting low free Al concentrations are not likely to affect plant growth, although the concentrations of potentially toxic Al3+ increased with soil depth due to higher total Al and lower dissolved organic matter concentrations in soil solutions. The Ca additions caused an increase of Al in the organic layer leachate, probably because Al3+ was exchanged against the added Ca2+ ions while pH remained constant. The free ion molar ratios of Ca2+:Al3+ (mean ratio ca. 400) were far above the threshold (≤ 1) for Al toxicity, because of a much higher degree of organo-complexation of Al than Ca. High Al fluxes in litterfall (8.8 – 14.2 kg ha−1 yr−1) indicate a high Al circulation through the ecosystem. The Al concentrations in the organic layer leachate were driven by the acidification of the ecosystem and increased significantly between 1999 and 2008. However, the Ca:Al molar ratios in organic layer leachate and all aboveground ecosystem solutions were above the threshold for Al toxicity. Except for two Al accumulating and one non-accumulating tree species, the Ca:Al molar ratios in tree leaves from the study area were above the Al toxicity threshold of 12.5. I conclude that toxic effects in the hydroponic experiment occurred at Al concentrations far above those in native organic layer leachate, shoot biomass production was likely inhibited by reduced Mg uptake, impairing photosynthesis, and the stimulation of root growth at low Al concentrations can be possibly attributed to improved P uptake. Dissolved organic matter in soil solutions detoxifies Al in acidic tropical forest soils and a wide distribution of Al accumulating tree species and high Al fluxes in the ecosystem do not necessarily imply a general Al phytotoxicity.

Long distance root–shoot signalling in plant–insect community interactions

Plants mediate interactions between insects, including leaf- and root-feeders; yet the underlying mechanisms and connection with ecological theory remain unresolved. In this review, based on novel insights into long-distance (i.e., leaf–leaf, root–shoot) defence signalling, we explore the role of phytohormones in driving broad-scale patterns of aboveground–belowground interactions that can be extrapolated to general plant–insect relationships. We propose that the outcome of intra-feeding guild interactions is generally negative due to induction of similar phytohormonal pathways, whereas between-guild interactions are often positive due to negative signal crosstalk. However, not all outcomes could be explained by feeding guild; we argue that future studies should target ecologically representative plant–insect systems, distinguish subguilds, and include plant growth hormones to improve our understanding of plant-mediated interactions.

Defence on demand: mechanisms behind optimal defence patterns

Background The optimal defence hypothesis (ODH) predicts that tissues that contribute most to a plant's fitness and have the highest probability of being attacked will be the parts best defended against biotic threats, including herbivores. In general, young sink tissues and reproductive structures show stronger induced defence responses after attack from pathogens and herbivores and contain higher basal levels of specialized defensive metabolites than other plant parts. However, the underlying physiological mechanisms responsible for these developmentally regulated defence patterns remain unknown. Scope This review summarizes current knowledge about optimal defence patterns in above- and below-ground plant tissues, including information on basal and induced defence metabolite accumulation, defensive structures and their regulation by jasmonic acid (JA). Physiological regulations underlying developmental differences of tissues with contrasting defence patterns are highlighted, with a special focus on the role of classical plant growth hormones, including auxins, cytokinins, gibberellins and brassinosteroids, and their interactions with the JA pathway. By synthesizing recent findings about the dual roles of these growth hormones in plant development and defence responses, this review aims to provide a framework for new discoveries on the molecular basis of patterns predicted by the ODH. Conclusions Almost four decades after its formulation, we are just beginning to understand the underlying molecular mechanisms responsible for the patterns of defence allocation predicted by the ODH. A requirement for future advances will be to understand how developmental and defence processes are integrated.

The Rice Transcription Factor WRKY53 Suppresses Herbivore-Induced Defenses by Acting as a Negative Feedback Modulator of Mitogen-Activated Protein Kinase Activity

The mechanisms by which herbivore-attacked plants activate their defenses are well studied. By contrast, little is known about the regulatory mechanisms that allow them to control their defensive investment and avoid a defensive overshoot. We characterized a rice (Oryza sativa) WRKY gene, OsWRKY53, whose expression is rapidly induced upon wounding and induced in a delayed fashion upon attack by the striped stem borer (SSB) Chilo suppressalis. The transcript levels of OsWRKY53 are independent of endogenous jasmonic acid but positively regulated by the mitogen-activated protein kinases OsMPK3/OsMPK6. OsWRKY53 physically interacts with OsMPK3/OsMPK6 and suppresses their activity in vitro. By consequence, it modulates the expression of defensive, MPK-regulated WRKYs and thereby reduces jasmonic acid, jasmonoyl-isoleucine, and ethylene induction. This phytohormonal reconfiguration is associated with a reduction in trypsin protease inhibitor activity and improved SSB performance. OsWRKY53 is also shown to be a negative regulator of plant growth. Taken together, these results show that OsWRKY53 functions as a negative feedback modulator of MPK3/MPK6 and thereby acts as an early suppressor of induced defenses. OsWRKY53 therefore enables rice plants to control the magnitude of their defensive investment during early signaling.

A fungal endophyte helps plants to tolerate root herbivory through changes in gibberellin and jasmonate signaling

Plant–microbe mutualisms can improve plant defense, but the impact of root endophytes on below-ground herbivore interactions remains unknown. We investigated the effects of the root endophyte Piriformospora indica on interactions between rice (Oryza sativa) plants and its root herbivore rice water weevil (RWW; Lissorhoptrus oryzophilus), and how plant jasmonic acid (JA) and GA regulate this tripartite interaction. Glasshouse experiments with wild-type rice and coi1-18 and Eui1-OX mutants combined with nutrient, jasmonate and gene expression analyses were used to test: whether RWW adult herbivory above ground influences subsequent damage caused by larval herbivory below ground; whether P. indica protects plants against RWW; and whether GA and JA signaling mediate these interactions. The endophyte induced plant tolerance to root herbivory. RWW adults and larvae acted synergistically via JA signaling to reduce root growth, while endophyte-elicited GA biosynthesis suppressed the herbivore-induced JA in roots and recovered plant growth. Our study shows for the first time the impact of a root endophyte on plant defense against below-ground herbivores, adds to growing evidence that induced tolerance may be an important root defense, and implicates GA as a signal component of inducible plant tolerance against biotic stress.

Establecimiento y desarrollo en el cultivo forzado de tomate : efecto de fitorreguladores

El proceso de fructificación y desarrollo en tomate puede ser inducido naturalmente por polinización o partenocarpia y artificialmente por aplicación de reguladores; esta respuesta es variable según tipo y dosis de hormona, momento de aplicación y cultivar involucrado. El objetivo de este trabajo fue evaluar la capacidad de auxinas y giberelinas para inducir el desarrollo partenocárpico en genotipos de crecimiento indeterminado. Como factores se consideraron tipo de regulador -AG3 y β-NOA- en dosis fija, momento de aplicación -0, 5, 12, 19, 26 dpa- y genotipo -Rutgers, Fortaleza F1 y Colt 45-. Las mejores respuestas a nivel de porcentaje de fructificación y peso fresco se obtuvieron con β-NOA en comparación con AG3. Considerando todos los factores analizados, solamente la aplicación de β-NOA a 5 dpa permitió alcanzar porcentajes de fructificación y tamaño final de frutos similares a los obtenidos por autopolinización. El período de sensibilidad y el tamaño final de los frutos presentaron interacción con las variables genotipo, momento de aplicación y tipo de regulador. Se observó además que AG3 provocó un escaso desarrollo placentario y ausencia de óvulos mientras que β-NOA indujo un desarrollo de placentas y óvulos similar al de los frutos obtenidos por autopolinización.

Establecimiento y desarrollo en el cultivo forzado de tomate : aplicación de dosis variables de fitorreguladores.

Las hormonas vegetales son capaces de controlar el desarrollo reproductivo, desde la diferenciación floral hasta los últimos estadios del desarrollo de los frutos. En particular, la etapa de fructificación y desarrollo depende del contenido endógeno de estas sustancias, y es posible manipular la iniciación del desarrollo del fruto por aplicación externa de hormonas. Previamente se evaluó el proceso de fructificación y desarrollo en el cultivo de tomate en invernadero en respuesta a la aplicación de b-NOA y AG3 en dosis fijas: se observó sensibilidad diferencial dependiendo del genotipo y tipo de regulador. El objetivo de este trabajo fue establecer dosis y momento óptimo para la aplicación de b-NOA y AG3 como formas de mejorar la fructificación y el desarrollo de frutos partenocárpicos. Como factores se consideraron tipo de regulador -b-NOA y AG3- en dosis y momentos de aplicación variables. Empleando ovarios no polinizados como sistema experimental fue posible concluir que la aplicación de 40 ppm de b-NOA a 7 días post antesis ofrece las mayores ventajas desde el punto de vista del rendimiento y menor impacto fisiológico, sin alterar el período de desarrollo de los frutos.

Fertilización del kiwi (Actinidia deliciosa (A. Chev.) C. F. Liang & A. R. Ferguson, cv. Hayward) durante la etapa de implantación en el sudeste bonaerense (Argentina)

En una plantación de kiwi situada en Balcarce (provincia de Buenos Aires, Argentina), se evaluó el efecto de la fertilización nitrogenada y potásica sobre el crecimiento de la planta durante la etapa de implantación, en la temporada 2008 - 2009. Los tratamientos fueron T: control sin fertilizar, N: fertilizado con nitrógeno (48 kg/ha N), K: fertilizado con potasio (60 kg/ha K) y NK: fertilizado con nitrógeno y potasio (48 kg/ha N - 60 kg/ha K). Se realizaron muestreos periódicos de: longitud de tallo, longitud del último entrenudo y área foliar. Se determinó el diámetro del tallo a fin de temporada y la distribución de raíces en el perfil de suelo en plena dormición. Se efectuaron análisis foliares de macronutrientes, así como N mineral, N anaeróbico (NAN) y carbono orgánico (CO) en el suelo. El suelo correspondió a un Argiudol, con niveles de NAN entre 85 - 100 mg/kg y de CO de 45 - 50 g/kg en los primeros 30 cm. Los resultados obtenidos, en cuanto al crecimiento de la parte aérea de la planta, no constituyen evidencia suficiente para justificar la fertilización con N y K bajo las condiciones experimentales del estudio. Aun en el T, los niveles foliares de N y K se mantuvieron dentro de los estándares normales. El sistema radical, observado durante el reposo, se habría visto beneficiado por la fertilización nitrogenada. En el estrato de suelo comprendido entre 10 y 20 cm se determinó un promedio de 61 raíces/100 cm2, en el tratamiento N, 40 en el tratamiento combinado NK, 28 en el tratamiento K, y 19 en el testigo.

Mineralogical-sedimentological and chemical investigation of sediments from Cross Bank off Florida

The sediments of a core of.1.55 m length taken on the windward side of the Cross Bank, Florida Bay, are clearly subdivided into two portions, as shown by grain size analysis: silt-sized particles predominate in the relatively homogeneous lower two thirds of the core. This is succeeded abruptly by a thin layer of sand, containing fragments of Halimeda. They indicate a catastrophic event in the Florida Bay region, because Halimeda does not grow within Florida Bay. Above this layer, the amount of sand decreases at first and then continuously increases right to the present sediment-water-interface. The median and skewness increase simultaneously with the increase in the sand and granule portion. We assume that the changing grain size distribution was determined chiefly by the density of the marine flora: during the deposition of the lower two thirds of the core a dense grass cover acted as a sediment catcher for the fine-grained detritus washed out of the shallow basins of the Florida Bay, and simultaneously prohibited renewed reworking. Similar processes go on today on the surface of most mud banks of Florida Bay. The catastrophic event indicated by the sand layer probably changed the morphology of the bank to such an extent that the sampling point was shifted more to the windward side of the bank. This side is characterized by less dense plant growth. Therefore, less detritus could be caught and the material deposited could be reworked. The pronounced increase in skewness in the upper third of the core certainly indicates a strong washing out of the smaller-sized particles. The sediments are predominantly made up of carbonates, averagely 88.14 percent. The average CaCO3-content is 83.87 percent and the average MgCO3-content amounts to 4.27 percent. The chief carbonate mineral is aragonite making up 60.1 percent of the carbonate portion in the average, followed by high-magnesian calcite (33.8 percent) and calcite (6.1 percent). With increasing grain size the aragonite clearly increases at the cost of high-magnesian calcite in the upper third of the core. Chemically, this is shown by an increase of the CaCO3 : MgCO3-ratio. This increase is mainly caused by the more common occurrence of aragonitic fragments of mollusks in the coarse grain fractions. The bulk of the carbonates is made up of mollusks, foraminifera, ostracods, and - to a much lesser extent - of corals, worm-tubes, coccolithophorids, and calcareous algae, as shown by microscopic investigations. The total amount of the carbonate in the sediments is biogenic detritus with the possible exception of a very small amount of aragonite needles in the clay and fine silt fraction. The individual carbonate components of the gravel and sand fraction can be relatively easy identified as members of a particular animal or plant group. This becomes very difficult in the silt and clay fraction. Brownish aggregates are very common in the coarse and medium silt fraction. It was not always possible to clarify their origin (biogenic detritus, faecal pellets or carbonate particles cemented by carbonates or organic slime, etc.). Organic matter (plant fragments, rootlets), quartz, opal (siliceous sponge needles), and feldspar also occur in the sediments, besides carbonates. The lowermost part of the core has an age of 1365 +/- 90 years, as shown by 14C analysis.

Seed Dispersal Spectrum of Woody Species in South Ecuadorian Dry Forests: Environmental Correlates and the Effect of Considering Species Abundance

This study examines the seed dispersal spectrum of the tropical dry forests of Southern Ecuador, in an effort to contribute to the knowledge of the complex dynamics of tropical dry forests. Seed dispersal spectrum was described for a total number of 160 species. Relationships of dispersal syndromes with plant growth form and climatic seasonality were explored. For a subset of 97 species, we determined whether dispersal spectrum changes when species abundance, in addition to species number, is taken into account. The same subset was used to relate dispersal syndromes with the environmental conditions. Zoochorous species dominated in the studied community. When considering the individual abundance of each species, however, anemochory was the prevalent dispersal syndrome. We found a significant difference in the frequency of dispersal syndromes among plant growth forms, with epizoochory only occurring in shrub species. The dispersal spectrum was dependent on climatic seasonality. The largest proportion of anemochorous species fructified during the dry season, while zoochorous diaspores dominated during the rainy season. A fourth-corner analysis indicated that the seed dispersal spectrum of Southern Ecuador dry forests is controlled by environmental conditions such as annual precipitation, annual temperature range or topography. Our results suggest that spatio-temporal changes in the environmental conditions may affect important ecological processes for dispersal. Thus, the predominance of one syndrome or another may depend on the spatial variation of environmental conditions.

Arabidopsis thaliana DOF6 negatively affects germination in non-after ripened seeds and interacts with TCP14

Seed dormancy prevents seeds from germinating under environmental conditions unfavourable for plant growth and development and constitutes an evolutionary advantage. Dry storage, also known as after-ripening, gradually decreases seed dormancy by mechanisms not well understood. An Arabidopsis thaliana DOF transcription factor gene (DOF6) affecting seed germination has been characterized. The transcript levels of this gene accumulate in dry seeds and decay gradually during after-ripening and also upon seed imbibition. While constitutive over-expression of DOF6 produced aberrant growth and sterility in the plant, its over-expression induced upon seed imbibition triggered delayed germination, abscisic acid (ABA)-hypersensitive phenotypes and increased expression of the ABA biosynthetic gene ABA1 and ABA-related stress genes. Wild-type germination and gene expression were gradually restored during seed after-ripening, despite of DOF6-induced over-expression. DOF6 was found to interact in a yeast two-hybrid system andin planta with TCP14, a previously described positive regulator of seed germination. The expression of ABA1 and ABA-related stress genes was also enhanced in tcp14 knock-out mutants. Taken together, these results indicate that DOF6 negatively affects seed germination and opposes TCP14 function in the regulation of a specific set of ABA-related genes

Aquaponics: integrating fish feeding rates and ion waste production for strawberry hydroponics

Aquaponics is the science of integrating intensive fish aquaculture with plant production in recirculating water systems. Although ion waste production by fish cannot satisfy all plant requirements, less is known about the relationship between total feed provided for fish and the production of milliequivalents (mEq) of different macronutrients for plants, especially for nutrient flow hydroponics used for strawberry production in Spain. That knowledge is essential to consider the amount of macronutrients available in aquaculture systems so that farmers can estimate how much nutrient needs to be supplemented in the waste water from fish, to produce viable plant growth. In the present experiment, tilapia (Oreochromis niloticus L.) were grown in a small-scale recirculating system at two different densities while growth and feed consumption were noted every week for five weeks. At the same time points, water samples were taken to measure pH, EC25, HCO3 – , Cl – , NH4 + , NO2 – , NO3 – , H2PO4 – , SO4 2– , Na + , K+ , Ca 2+ and Mg 2+ build up. The total increase in mEq of each ion per kg of feed provided to the fish was highest for NO3 - , followed, in decreasing order, by Ca 2+ , H2PO4 – , K+ , Mg 2+ and SO4 2– . The total amount of feed required per mEq ranged from 1.61- 13.1 kg for the four most abundant ions (NO3 – , Ca 2+ , H2PO4 – and K+ ) at a density of 2 kg fish m–3 , suggesting that it would be rather easy to maintain small populations of fish to reduce the cost of hydroponic solution supplementation for strawberries.