Etiology and epidemiology of Pythium root rot in hydroponic crops: current knowledge and perspectives

The etiology and epidemiology of Pythium root rot in hydroponically-grown crops are reviewed with emphasis on knowledge and concepts considered important for managing the disease in commercial greenhouses. Pythium root rot continually threatens the productivity of numerous kinds of crops in hydroponic systems around the world including cucumber, tomato, sweet pepper, spinach, lettuce, nasturtium, arugula, rose, and chrysanthemum. Principal causal agents include Pythium aphanidermatum, Pythium dissotocum, members of Pythium group F, and Pythium ultimum var. ultimum. Perspectives are given of sources of initial inoculum of Pythium spp. in hydroponic systems, of infection and colonization of roots by the pathogens, symptom development and inoculum production in host roots, and inoculum dispersal in nutrient solutions. Recent findings that a specific elicitor produced by P. aphanidermatum may trigger necrosis (browning) of the roots and the transition from biotrophic to necrotrophic infection are considered. Effects on root rot epidemics of host factors (disease susceptibility, phenological growth stage, root exudates and phenolic substances), the root environment (rooting media, concentrations of dissolved oxygen and phenolic substances in the nutrient solution, microbial communities and temperature) and human interferences (cropping practices and control measures) are reviewed. Recent findings on predisposition of roots to Pythium attack by environmental stress factors are highlighted. The commonly minor impact on epidemics of measures to disinfest nutrient solution as it recirculates outside the crop is contrasted with the impact of treatments that suppress Pythium in the roots and root zone of the crop. New discoveries that infection of roots by P. aphanidermatum markedly slows the increase in leaf area and whole-plant carbon gain without significant effect on the efficiency of photosynthesis per unit area of leaf are noted. The platform of knowledge and understanding of the etiology and epidemiology of root rot, and its effects on the physiology of the whole plant, are discussed in relation to new research directions and development of better practices to manage the disease in hydroponic crops. Focus is on methods and technologies for tracking Pythium and root rot, and on developing, integrating, and optimizing treatments to suppress the pathogen in the root zone and progress of root rot.

Uneven distribution of nutrients in the root zone affects the incidence of blossom end rot and concentration of calcium and potassium in fruits of tomato

Tomato plants ( Lycopersicon esculentum Mill. var. DRK) were grown hydroponically to determine the effect of an uneven distribution of nutrients in the root zone on blossomend rot (BER) and Ca and K concentrations in the fruits. The plants were grown in rockwool with their root system divided into two portions. Each portion was irrigated with nutrient solutions with either the same or the different electrical conductivity (EC) in the range 0 to 6 dS m(-1). Solutions with high EC supplied to both sides of the root system significantly increased the incidence of BER. However, when only water or a solution of low EC was supplied to one portion, BER was reduced by 80%. Fruit yields were significantly higher ( P < 0.01) for plants that received solutions of the uneven EC treatments (6/0 or 4.5/0 EC treatment). Plants supplied with solutions of uneven EC generally had higher leaf and fruit concentrations of Ca but lower concentrations of K than those supplied with solutions of high EC. There was no difference in Ca concentration at the distal end of young fruits of the uneven EC treatment but it was reduced in the high EC treatments. The concentration of K in the mature fruits of the uneven EC treatments was lower than that of the high EC treatments and higher or similar that of the 3/3 or 2.5/2.5 EC treatments ( controls). A clear relationship was found between the incidence of BER and the exudation rate. High rate of xylem exudation was observed in the uneven EC treatments. Reduction of BER in the uneven EC treatments is most likely to be the effect of high exudation rate on Ca status in the young fruits. It was concluded that high EC of solution had positive effects on Ca concentration and incidence of BER provided that nutrient solution with low EC or water is supplied to the one portion of the root system.

Root pruning as a means to encourage root growth in two ornamental shrubs, Buddleja davidii ‘Summer Beauty’ and Cistus ‘Snow Fire'

The inability of a plant to grow roots rapidly upon transplanting is one of the main factors contributing to poor establishment. In bare-rooted trees, treatments such as root pruning or application of the plant hormone auxin [e.g., indole butyric acid (IBA)] can promote root growth and aid long-term establishment. There is little information on ornamental containerised plants, however, other than the anecdotal notion that 'teasing' the roots out of the rootsoil mass before transplanting can be beneficial. In the present study we tested the ability of various root-pruning treatments and application of IBA to encourage new root and shoot growth in two shrub species, commonly produced in containers - Buddleja davidii 'Summer Beauty' and Cistus 'Snow Fire'. In a number of experiments, young plants were exposed to root manipulation (teasing, light pruning, or two types of heavy pruning) and/or treatment with IBA (at 500 or 1,000 mg l-1) before being transplanted into larger containers containing a medium of 1:1:1 (v/v/v) fine bark, sand and loam. Leaf stomatal conductance (gl) was measured 20 min, and 1, 2, 4 and 6 h after root manipulation. Net leaf CO2 assimilation (A) was measured frequently during the first week after transplanting, then at regular intervals up to 8 weeks after transplanting. Plants were harvested 8 weeks after transplanting, and root and shoot weights were measured. In both species, light root pruning alone, or in combination with 500 mg l-1 IBA, was most effective in stimulating root growth. In contrast, teasing, which is commonly used, showed no positive effect on root growth in Buddleja, and decreased new root growth in Cistus. The requirement for exogenous auxin to encourage new root growth varied between experiments and appeared to be influenced by the age and developmental stage of the plants. There were no consistent responses between root treatments and net CO2 assimilation rates, and changes in root weight were not closely correlated with changes in assimilation. The mechanisms whereby new root growth is sustained are discussed.

Uneven distribution of nutrients in the root zone affects the incidence of blossom end rot and concentration of calcium and potassium in fruits of tomato

Tomato plants ( Lycopersicon esculentum Mill. var. DRK) were grown hydroponically to determine the effect of an uneven distribution of nutrients in the root zone on blossomend rot (BER) and Ca and K concentrations in the fruits. The plants were grown in rockwool with their root system divided into two portions. Each portion was irrigated with nutrient solutions with either the same or the different electrical conductivity (EC) in the range 0 to 6 dS m(-1). Solutions with high EC supplied to both sides of the root system significantly increased the incidence of BER. However, when only water or a solution of low EC was supplied to one portion, BER was reduced by 80%. Fruit yields were significantly higher ( P < 0.01) for plants that received solutions of the uneven EC treatments (6/0 or 4.5/0 EC treatment). Plants supplied with solutions of uneven EC generally had higher leaf and fruit concentrations of Ca but lower concentrations of K than those supplied with solutions of high EC. There was no difference in Ca concentration at the distal end of young fruits of the uneven EC treatment but it was reduced in the high EC treatments. The concentration of K in the mature fruits of the uneven EC treatments was lower than that of the high EC treatments and higher or similar that of the 3/3 or 2.5/2.5 EC treatments ( controls). A clear relationship was found between the incidence of BER and the exudation rate. High rate of xylem exudation was observed in the uneven EC treatments. Reduction of BER in the uneven EC treatments is most likely to be the effect of high exudation rate on Ca status in the young fruits. It was concluded that high EC of solution had positive effects on Ca concentration and incidence of BER provided that nutrient solution with low EC or water is supplied to the one portion of the root system.

Elevated atmospheric carbon dioxide impairs the performance of root-feeding vine weevils by modifying root growth and secondary metabolites

Predicting how insect crop pests will respond to global climate change is an important part of increasing crop production for future food security, and will increasingly rely on empirically based evidence. The effects of atmospheric composition, especially elevated carbon dioxide (eCO(2)), on insect herbivores have been well studied, but this research has focussed almost exclusively on aboveground insects. However, responses of root-feeding insects to eCO(2) are unlikely to mirror these trends because of fundamental differences between aboveground and belowground habitats. Moreover, changes in secondary metabolites and defensive responses to insect attack under eCO(2) conditions are largely unexplored for root herbivore interactions. This study investigated how eCO(2) (700 mu mol mol-1) affected a root-feeding herbivore via changes to plant growth and concentrations of carbon (C), nitrogen (N) and phenolics. This study used the root-feeding vine weevil, Otiorhynchus sulcatus and the perennial crop, Ribes nigrum. Weevil populations decreased by 33% and body mass decreased by 23% (from 7.2 to 5.4 mg) in eCO(2). Root biomass decreased by 16% in eCO(2), which was strongly correlated with weevil performance. While root N concentrations fell by 8%, there were no significant effects of eCO(2) on root C and N concentrations. Weevils caused a sink in plants, resulting in 8-12% decreases in leaf C concentration following herbivory. There was an interactive effect of CO(2) and root herbivory on root phenolic concentrations, whereby weevils induced an increase at ambient CO(2), suggestive of defensive response, but caused a decrease under eCO(2). Contrary to predictions, there was a positive relationship between root phenolics and weevil performance. We conclude that impaired root-growth underpinned the negative effects of eCO(2) on vine weevils and speculate that the plant's failure to mount a defensive response at eCO(2) may have intensified these negative effects.

A tillering inhibition gene influences root-shoot carbon partitioning and pattern of water use to improve wheat productivity in rainfed environments

Genetic modification of shoot and root morphology has potential to improve water and nutrient 19 uptake of wheat crops in rainfed environments. Near-isogenic lines (NILs) varying for a tillering 20 inhibition (tin) gene and representing multiple genetic backgrounds were investigated in contrasting 21 controlled environments for shoot and root growth. Leaf area, shoot and root biomass were similar 22 until tillering whereupon reduced tillering in tin-containing NILs produced reductions of up to 60% in 23 total leaf area and biomass, and increases in total root length of up to 120% and root biomass to 24 145%. Together, root-to-shoot ratio increased two-fold with the tin gene. The influence of tin on shoot 25 and root growth was greatest in the cv. Banks genetic background, particularly in the biculm-selected 26 NIL, and was typically strongest in cooler environments. A separate de-tillering study confirmed 27 greater root-to-shoot ratios with regular tiller removal in non-tin containing genotypes. In validating 28 these observations in a rainfed field study, the tin allele had a negligible effect on seedling growth but 29 was associated with significantly (P<0.05) reduced tiller number (-37%), leaf area index (-26%) and 30 spike number (-35%) to reduce plant biomass (-19%) at anthesis. Root biomass, root-to-shoot ratio at 31 early stem elongation and root depth at maturity were increased in tin-containing NILs. Soil water use 32 was slowed in tin-containing NILs resulting in greater water availability, greater stomatal 33 conductance, cooler canopy temperatures and maintenance of green leaf area during grain-filling. 34 Together these effects contributed to increases in harvest index and grain yield. In both the controlled 35 and field environments, the tin gene was commonly associated with increased root length and biomass 36 but the significant influence of genetic background and environment suggests careful assessment of 37 tin-containing progeny in selection for genotypic increases in root growth.

Cotton root and shoot growth as affected by application of mepiquat chloride to cotton seeds

Estudou-se o efeito do tratamento de sementes de algodão com cloreto de mepiquat sobre o crescimento inicial de raízes e parte aérea. O experimento, realizado em casa de vegetação, utilizou vasos de PVC adaptados com uma parede frontal de vidro e os tratamentos foram constituídos por cinco doses do cloreto de mepiquat (CM) do ingrediente ativo (i.a.): 0, 3, 6, 9 e 12 g kg-1 de sementes, pulverizado sobre as sementes, e a cultivar FM 993. Massa de matéria seca da parte aérea (folhas, pecíolos e haste), massa de matéria seca da raiz, área foliar, relação parte aérea:raiz, relação área foliar:crescimento radicular, o comprimento da parte aérea foram avaliados aos 21 dias após a semeadura. Crescimento radicular foi avaliado a cada três dias até os 18 dias. O CM aplicado às sementes do algodão promove redução da altura da planta e da área foliar, sem, contudo, afetar produção de massa de matéria seca da parte aérea e raiz, relação parte aérea:raiz, relação área foliar:crescimento radicular e comprimento total de raízes do algodoeiro. Assim, no presente experimento não foi observado efeito negativo do CM aplicado às sementes do algodoeiro na absorção de água pela planta.

Developmental and structural features of secretory canals in root and shoot wood of Copaifera langsdorffii Desf. (Leguminosae-Caesalpinioideae)

Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP)

On the taxonomic value of the anatomical structure of vegetative organs and inflorescence axis of Abolboda species (Xyridaceae - Poales)

Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq)

Morphology and anatomy of the vegetative organs and scapes from Aphorocaulon (Paepalanthus, Eriocaulaceae)

Estudou-se a anatomia de raízes, caules, folhas e escapos de espécies de Paepalanthus subseção Aphorocaulon. Estas plantas apresentam caules reduzidos com folhas em roseta, de onde crescem os paracládios (sistemas de inflorescências). As espécies apresentam raízes com epiderme unisseriada e córtex com células isodiamétricas. Tanto os caules reduzidos como os paracládios apresentam espessamento resultante da atividade do periciclo, denominado Meristema de Espessamento Primário (MEP). Ambos apresentam estrutura anatômica semelhante. Os escapos apresentam endoderme descontínua, periciclo sinuoso, o córtex apresenta costelas salientes (5-6). As folhas apresentam células epidérmicas alongadas no sentido longitudinal com paredes levemente espessadas, estômatos somente na face abaxial, com câmara subestomática especializada, feixes vasculares colaterais com bainha dupla. Essas estruturas anatômicas são comuns para as espécies da subseção Aphorocaulon. Algumas características anatômicas observadas nestas espécies são típicas de plantas que crescem nos campos rupestres.

Root canal treatment of three-rooted maxillary first and second premolars - a case report

Aim To report the diagnosis and successful clinical management of a patient with anatomical variation in maxillary first and second premolars.Summary Maxillary premolars have a highly variable root canal morphology, but it is rare, especially in the second premolar, to find three canals. This article describes the diagnosis and clinical management of first and second premolars with three canals and three separate roots, drawing particular attention to radiographic interpretation and access refinements.

Mixing Eucalyptus and Acacia trees leads to fine root over-yielding and vertical segregation between species

The consequences of diversity on belowground processes are still poorly known in tropical forests. The distributions of very fine roots (diameter <1 mm) and fine roots (diameter <3 mm) were studied in a randomized block design close to the harvest age of fast-growing plantations. A replacement series was set up in Brazil with mono-specific Eucalyptus grandis (100E) and Acacia mangium (100A) stands and a mixture with the same stocking density and 50 % of each species (50A:50E). The total fine root (FR) biomass down to a depth of 2 m was about 27 % higher in 50A:50E than in 100A and 100E. Fine root over-yielding in 50A:50E resulted from a 72 % rise in E. grandis fine root biomass per tree relative to 100E, whereas A. mangium FR biomass per tree was 17 % lower than in 100A. Mixing A. mangium with E. grandis trees led to a drop in A. mangium FR biomass in the upper 50 cm of soil relative to 100A, partially balanced by a rise in deep soil layers. Our results highlight similarities in the effects of directional resources on leaf and FR distributions in the mixture, with A. mangium leaves below the E. grandis canopy and a low density of A. mangium fine roots in the resource-rich soil layers relative to monospecific stands. The vertical segregation of resource-absorbing organs did not lead to niche complementarity expected to increase the total biomass production. © 2012 Springer-Verlag Berlin Heidelberg.

Organic fertilization of peach trees: implication on nitrogen availability, root growth and carbon distribution within plant

In the last years, sustainable horticulture has been increasing; however, to be successful this practice needs an efficient soil fertility management to maintain a high productivity and fruit quality standards. For this purpose composted organic materials from agri-food industry and municipal solid waste has been used as a source to replace chemical fertilizers and increase soil organic matter. To better understand the influence of compost application on soil fertility and plant growth, we carried out a study comparing organic and mineral nitrogen (N) fertilization in micro propagated plants, potted trees and commercial peach orchard with these aims: 1. evaluation of tree development, CO2 fixation and carbon partition to the different organs of two-years-old potted peach trees. 2. Determination of soil N concentration and nitrate-N effect on plant growth and root oxidative stress of micro propagated plant after increasing rates of N applications. 3. Assessment of soil chemical and biological fertility, tree growth and yield and fruit quality in a commercial orchard. The addition of compost at high rate was effective in increasing CO2 fixation, promoting root growth, shoot and fruit biomass. Furthermore, organic fertilizers influenced C partitioning, favoring C accumulation in roots, wood and fruits. The higher CO2 fixation was the result of a larger tree leaf area, rather than an increase in leaf photosynthetic efficiency, showing a stimulation of plant growth by application of compost. High concentrations of compost increased total soil N concentration, but were not effective in increasing nitrate-N soil concentration; in contrast mineral-N applications increased linearly soil nitrate-N, even at the lowest rate tested. Soil nitrate-N concentration influenced positively plant growth at low rate (60- 80 mg kg-1), whereas at high concentrations showed negative effects. In this trial, the decrease of root growth, as a response to excessive nitrate-N soil concentration, was not anticipated by root oxidative stress. Continuous annual applications of compost for 10 years enhanced soil organic matter content and total soil N concentration. Additionally, high rate of compost application (10 t ha-1 year-1) enhanced microbial biomass. On the other hand, different fertilizers management did not modify tree yield, but influenced fruit size and precocity index. The present data support the idea that organic fertilizers can be used successfully as a substitute of mineral fertilizers in fruit tree nutrient management, since they promote an increase of soil chemical and biological fertility, prevent excessive nitrate-N soil concentration, promote plant growth and potentially C sequestration into the soil.

Factors controlling decomposition rates of fine root litter in temperate forests and grasslands

Background and aims Fine root decomposition contributes significantly to element cycling in terrestrial ecosystems. However, studies on root decomposition rates and on the factors that potentially influence them are fewer than those on leaf litter decomposition. To study the effects of region and land use intensity on fine root decomposition, we established a large scale study in three German regions with different climate regimes and soil properties. Methods In 150 forest and 150 grassland sites we deployed litterbags (100 μm mesh size) with standardized litter consisting of fine roots from European beech in forests and from a lowland mesophilous hay meadow in grasslands. In the central study region, we compared decomposition rates of this standardized litter with root litter collected on-site to separate the effect of litter quality from environmental factors. Results Standardized herbaceous roots in grassland soils decomposed on average significantly faster (24 ± 6 % mass loss after 12 months, mean ± SD) than beech roots in forest soils (12 ± 4 %; p < 0.001). Fine root decomposition varied among the three study regions. Land use intensity, in particular N addition, decreased fine root decomposition in grasslands. The initial lignin:N ratio explained 15 % of the variance in grasslands and 11 % in forests. Soil moisture, soil temperature, and C:N ratios of soils together explained 34 % of the variance of the fine root mass loss in grasslands, and 24 % in forests. Conclusions Grasslands, which have higher fine root biomass and root turnover compared to forests, also have higher rates of root decomposition. Our results further show that at the regional scale fine root decomposition is influenced by environmental variables such as soil moisture, soil temperature and soil nutrient content. Additional variation is explained by root litter quality.

Induced carbon reallocation and compensatory growth as root herbivore tolerance mechanisms

Upon attack by leaf herbivores, many plants reallocate photoassimilates below ground. However, little is known about how plants respond when the roots themselves come under attack. We investigated induced resource allocation in maize plants that are infested by the larvae Western corn rootworm Diabrotica virgifera virgifera. Using radioactive 11CO2, we demonstrate that root-attacked maize plants allocate more new 11C carbon from source leaves to stems, but not to roots. Reduced meristematic activity and reduced invertase activity in attacked maize root systems are identified as possible drivers of this shoot reallocation response. The increased allocation of photoassimilates to stems is shown to be associated with a marked thickening of these tissues and increased growth of stem-borne crown roots. A strong quantitative correlation between stem thickness and root regrowth across different watering levels suggests that retaining photoassimilates in the shoots may help root-attacked plants to compensate for the loss of belowground tissues. Taken together, our results indicate that induced tolerance may be an important strategy of plants to withstand belowground attack. Furthermore, root herbivore-induced carbon reallocation needs to be taken into account when studying plant-mediated interactions between herbivores.