Morphological and physiological responses of rice seedlings to complete submergence (flash flooding)

Background and Aims

Reducing damage to rice seedlings caused by flash flooding will improve the productivity of rainfed lowland rice in West Africa. Accordingly, the morphological and physiological responses of different forms of rice to complete submergence were examined in field and pot experiments to identify primary causes of damage.

Methods

To characterize the physiological responses, seedlings from a wide genetic base including Oryza sativa, O. glaberrima and interspecific hybrids were compared using principle component analysis.

Key Results

Important factors linked to flash-flood tolerance included minimal shoot elongation underwater, increase in dry matter weight during submergence and post-submergence resistance to lodging. In particular, fast shoot elongation during submergence negatively affected plant growth after de-submergence. Also shoot-elongating cultivars showed a strong negative correlation between dry matter weight of the leaves that developed before submergence and leaves developing during submergence.

Conclusions

Enhancement of shoot elongation during submergence in water that is too deep to permit re-emergence by small seedlings represents a futile escape strategy that takes place at the expense of existing dry matter in circumstances where underwater photosynthetic carbon fixation is negligible. Consequently, it compromises survival or recovery growth once flood water levels recede and plants are re-exposed to the aerial environment. Tolerance is greater in cultivars where acceleration of elongation caused by submergence is minimal.Key words: Africa, flash floods, Oryza glaberrima, rainfed lowland, rice, shoot elongation, stress tolerance, submergence     

Is elongation-induced leaf emergence beneficial for submerged Rumex species?

Background and Aims

Plant species from various taxa ‘escape’ from low oxygen conditions associated with submergence by a suite of traits collectively called the low oxygen escape syndrome (LOES). The expression of these traits is associated with costs and benefits. Thus far, remarkably few studies have dealt with the expected benefits of the LOES.

Methods

Young plants were fully submerged at initial depths of 450 mm (deep) or 150–240 mm (shallow). Rumex palustris leaf tips emerged from the shallow flooding within a few days, whereas a slight lowering of shallow flooding was required to expose R. acetosa leaf tips to the atmosphere. Shoot biomass and petiole porosity were measured for all species, and treatments and data from the deep and shallow submergence treatments were compared with non-flooded controls.

Key Results

R. palustris is characterized by submergence-induced enhanced petiole elongation. R. acetosa lacked this growth response. Upon leaf tip emergence, R. palustris increased its biomass, whereas R. acetosa did not. Furthermore, petiole porosity in R. palustris was twice as high as in R. acetosa.

Conclusions

Leaf emergence restores gas exchange between roots and the atmosphere in R. palustris. This occurs to a much lesser extent in R. acetosa and is attributable to its lower petiole porosity and therefore limited internal gas transport. Leaf emergence resulting from fast petiole elongation appears to benefit biomass accumulation if these plants contain sufficient aerenchyma in petioles and roots to facilitate internal gas exchange.Key words: Submergence, emergence, enhanced shoot elongation, porosity, aerenchyma, Rumex, cost–benefit analysis, phenotypic plasticity     

Distinct mechanisms for aerenchyma formation in leaf sheaths of rice genotypes displaying a quiescence or escape strategy for flooding tolerance

Background and Aims

Rice is one of the few crops able to withstand periods of partial or even complete submergence. One of the adaptive traits of rice is the constitutive presence and further development of aerenchyma which enables oxygen to be transported to submerged organs. The development of lysigenous aerenchyma is promoted by ethylene accumulating within the submerged plant tissues, although other signalling mechanisms may also co-exist. In this study, aerenchyma development was analysed in two rice (Oryza sativa) varieties, ‘FR13A’ and ‘Arborio Precoce’, which show opposite traits in flooding response in terms of internode elongation and survival.

Methods

The growth and survival of rice varieties under submergence was investigated in the leaf sheath of ‘FR13A’ and ‘Arborio Precoce’. The possible involvement of ethylene and reactive oxygen species (ROS) was evaluated in relation to aerenchyma formation. Cell viability and DNA fragmentation were determined by FDA/FM4-64 staining and TUNEL assay, respectively. Ethylene production was monitored by gas chromatography and by analysing ACO gene expression. ROS production was measured by using Amplex Red assay kit and the fluorescent dye DCFH₂-DA. The expression of APX1 was also evaluated. AVG and DPI solutions were used to test the effect of inhibiting ethylene biosynthesis and ROS production, respectively.

Key Results

Both the varieties displayed constitutive lysigenous aerenchyma formation, which was further enhanced when submerged. ‘Arborio Precoce’, which is characterized by fast elongation when submerged, showed active ethylene biosynthetic machinery associated with increased aerenchymatous areas. ‘FR13A’, which harbours the Sub1A gene that limits growth during oxygen deprivation, did not show any increase in ethylene production after submersion but still displayed increased aerenchyma. Hydrogen peroxide levels increased in ‘FR13A’ but not in ‘Arborio Precoce’.

Conclusions

While ethylene controls aerenchyma formation in the fast-elongating ‘Arborio Precoce’ variety, in ‘FR13A’ ROS accumulation plays an important role.     

Recovery dynamics of growth, photosynthesis and carbohydrate accumulation after de-submergence: a comparison between two wetland plants showing escape and quiescence strategies

Background and Aims

The capacity for fast-growth recovery after de-submergence is important for establishment of riparian species in a water-level-fluctuation zone. Recovery patterns of two wetland plants, Alternanthera philoxeroides and Hemarthria altissima, showing ‘escape’ and ‘quiescence’ responses, respectively, during submergence were investigated.

Methods

Leaf and root growth and photosynthesis were monitored continuously during 10 d of recovery following 20 d of complete submergence. Above- and below-ground dry weights, as well as carbohydrate concentrations, were measured several times during the experiment.

Key Results

Both species remobilized stored carbohydrate during submergence. Although enhanced internode elongation depleted the carbohydrate storage in A. philoxeroides during submergence, this species resumed leaf growth 3 d after de-submergence concomitant with restoration of the maximal photosynthetic capacity. In contrast, some sucrose was conserved in shoots of H. altissima during submergence, which promoted rapid re-growth of leaves 2 d after de-submergence and earlier than the full recovery of photosynthesis. The recovery of root growth was delayed by 1–2 d compared with leaves in both species.

Conclusions

Submergence tolerance of the escape and quiescence strategies entails not only the corresponding regulation of growth, carbohydrate catabolism and energy metabolism during submergence but also co-ordinated recovery of photosynthesis, growth and carbohydrate partitioning following de-submergence.     

Escape from water or remain quiescent? Lotus tenuis changes its strategy depending on depth of submergence

Background and Aims

Two main strategies that allow plants to cope with soil waterlogging or deeper submergence are: (1) escaping by means of upward shoot elongation or (2) remaining quiescent underwater. This study investigates these strategies in Lotus tenuis, a forage legume of increasing importance in areas prone to soil waterlogging, shallow submergence or complete submergence.

Methods

Plants of L. tenuis were subjected for 30 d to well-drained (control), waterlogged (water-saturated soil), partially submerged (6 cm water depth) and completely submerged conditions. Plant responses assessed were tissue porosity, shoot number and length, biomass and utilization of water-soluble carbohydrates (WSCs) and starch in the crown.

Key Results

Lotus tenuis adjusted its strategy depending on the depth of submergence. Root growth of partially submerged plants ceased and carbon allocation prioritized shoot lengthening (32 cm vs. 24·5 cm under other treatments), without depleting carbohydrate reserves to sustain the faster growth. These plants also developed more shoot and root porosity. In contrast, completely submerged plants became quiescent, with no associated biomass accumulation, new shoot production or shoot elongation. In addition, tissue porosity was not enhanced. The survival of completely submerged plants is attributed to consumption of WSCs and starch reserves from crowns (concentrations 50–75 % less than in other treatments).

Conclusions

The forage legume L. tenuis has the flexibility either to escape from partial submergence by elongating its shoot more vigorously to avoid becoming totally submerged or to adopt a non-elongating quiescent strategy when completely immersed that is based on utilizing stored reserves. The possession of these alternative survival strategies helps to explain the success of L. tenuis in environments subjected to unpredictable flooding depths.     

Wait or escape? Contrasting submergence tolerance strategies of Rorippa amphibia,Rorippa sylvestris and their hybrid

Background and Aims

Differential responses of closely related species to submergence can provide insight into the evolution and mechanisms of submergence tolerance. Several traits of two wetland species from habitats with contrasting flooding regimes, Rorippa amphibia and Rorippa sylvestris, as well as F₁ hybrid Rorippa × anceps were analysed to unravel mechanisms underlying submergence tolerance.

Methods

In the first submergence experiment (lasting 20 d) we analysed biomass, stem elongation and carbohydrate content. In the second submergence experiment (lasting 3 months) we analysed survival and the effect of re-establishment of air contact on biomass and carbohydrate content. In a separate experiment we analysed expression of two carbohydrate catabolism genes, ADH1 and SUS1, upon re-establishment of air contact following submergence.

Key Results

All plants had low mortality even after 3 months of submergence. Rorippa sylvestris was characterized by 100 % survival and higher carbohydrate levels coupled with lower ADH1 gene expression as well as reduced growth compared with R. amphibia. Rorippa amphibia and the hybrid elongated their stems but this did not pay-off in higher survival when plants remained submerged. Only R. amphibia and the hybrid benefited in terms of increased biomass and carbohydrate accumulation upon re-establishing air contact.

Conclusions

Results demonstrate contrasting ‘escape’ and ‘quiescence’ strategies between Rorippa species. Being a close relative of arabidopsis, Rorippa is an excellent model for future studies on the molecular mechanism(s) controlling these strategies.     

A kinetic analysis of hyponastic growth and petiole elongation upon ethylene exposure in Rumex palustris

Background and Aims

Complete submergence is an important stress factor for many terrestrial plants, and a limited number of species have evolved mechanisms to deal with these conditions. Rumex palustris is one such species and manages to outgrow the water, and thus restore contact with the atmosphere, through upward leaf growth (hyponasty) followed by strongly enhanced petiole elongation. These responses are initiated by the gaseous plant hormone ethylene, which accumulates inside plants due to physical entrapment. This study aimed to investigate the kinetics of ethylene-induced leaf hyponasty and petiole elongation.

Methods

Leaf hyponasty and petiole elongation was studied using a computerized digital camera set-up followed by image analyses. Linear variable displacement transducers were used for fine resolution monitoring and measurement of petiole growth rates.

Key Results

We show that submergence-induced hyponastic growth and petiole elongation in R. palustris can be mimicked by exposing plants to ethylene. The petiole elongation response to ethylene is shown to depend on the initial angle of the petiole. When petiole angles were artificially kept at 0°, rather than the natural angle of 35°, ethylene could not induce enhanced petiole elongation. This is very similar to submergence studies and confirms the idea that there are endogenous, angle-dependent signals that influence the petiole elongation response to ethylene.

Conclusions

Our data suggest that submergence and ethylene-induced hyponastic growth and enhanced petiole elongation responses in R. palustris are largely similar. However, there are some differences that may relate to the complexity of the submergence treatment as compared with an ethylene treatment.     

Tolerance of combined submergence and salinity in the halophytic stem-succulent Tecticornia pergranulata

Background and Aims

Habitats occupied by many halophytes are not only saline, but are also prone to flooding. Few studies have evaluated submergence tolerance in halophytes.

Methods

Responses to submergence, at a range of salinity levels, were studied for the halophytic stem-succulent Tecticornia pergranulata subsp. pergranulata (syn. Halosarcia pergranulata subsp. pergranulata). Growth and total sugars in succulent stems were assessed as a function of time after submergence. Underwater net photosynthesis, dark respiration, total sugars, glycinebetaine, Na⁺, Cl⁻ and K⁺, in succulent stems, were assessed in a NaCl dose-response experiment.

Key Results

Submerged plants ceased to grow, and tissue sugars declined. Photosynthesis by succulent stems was reduced markedly when underwater, as compared with in air. Capacity for underwater net photosynthesis (P_N) was not affected by 10–400 mm NaCl, but it was reduced by 30 % at 800 mm. Dark respiration, underwater, increased in succulent stems at 200–800 mm NaCl, as compared with those at 10 mm NaCl. On an ethanol-insoluble dry mass basis, K⁺ concentration in succulent stems of submerged plants was equal to that in drained controls, across all NaCl treatments. Na⁺ and Cl⁻ concentrations, however, were elevated in stems of submerged plants, but so was glycinebetaine. Submerged stems increased in succulence, so solutes would have been ‘diluted’ on a tissue-water basis.

Conclusions

Tecticornia pergranulata tolerates complete submergence, even in waters of high salinity. A ‘quiescence response’, i.e. no shoot growth, would conserve carbohydrates, but tissue sugars still declined with time. A low K⁺ : Na⁺ ratio, typical for tissues of succulent halophytes, was tolerated even during prolonged submergence, as evidenced by maintenance of underwater P_N at up to 400 mm NaCl. Underwater P_N provides O₂ and sugars, and thus should enhance survival of submerged plants.Key words: Flooding, halophyte, Halosarcia pergranulata, inundation, inland salt marsh, respiration, Salicornioideae, salt lake, submergence–salinity interaction, tissue solutes, underwater net photosynthesis     

Intraspecific variation in the magnitude and pattern of flooding-induced shoot elongation in Rumex palustris

Background and Aims

Intraspecific variation in flooding tolerance is the basic pre-condition for adaptive flooding tolerance to evolve, and flooding-induced shoot elongation is an important trait that enables plants to survive shallow, prolonged flooding. Here an investigation was conducted to determine to what extent variation in flooding-induced leaf elongation exists among and within populations of the wetland species Rumex palustris, and whether the magnitude of elongation can be linked to habitat characteristics.

Methods

Offspring of eight genotypes collected in each of 12 populations from different sites (ranging from river mudflats with dynamic flooding regimes to areas with stagnant water) were submerged, and petioles, laminas and roots were harvested separately to measure traits related to elongation and plant growth.

Key Results

We found strong elongation of petioles upon submergence, and both among- and within-population variation in this trait, not only in final length, but also in the timing of the elongation response. However, the variation in elongation responses could not be linked to habitat type.

Conclusions

Spatio-temporal variation in the duration and depth of flooding in combination with a presumably weak selection against flooding-induced elongation may have contributed to the maintenance of large genetic variation in flooding-related traits among and within populations.     

Aquatic adventitious root development in partially and completely submerged wetland plants Cotula coronopifolia and Meionectes brownii

Background and Aims

A common response of wetland plants to flooding is the formation of aquatic adventitious roots. Observations of aquatic root growth are widespread; however, controlled studies of aquatic roots of terrestrial herbaceous species are scarce. Submergence tolerance and aquatic root growth and physiology were evaluated in two herbaceous, perennial wetland species Cotula coronopifolia and Meionectes brownii.

Methods

Plants were raised in large pots with ‘sediment’ roots in nutrient solution and then placed into individual tanks and shoots were left in air or submerged (completely or partially). The effects on growth of aquatic root removal, and of light availability to submerged plant organs, were evaluated. Responses of aquatic root porosity, chlorophyll and underwater photosynthesis, were studied.

Key Results

Both species tolerated 4 weeks of complete or partial submergence. Extensive, photosynthetically active, aquatic adventitious roots grew from submerged stems and contributed up to 90 % of the total root dry mass. When aquatic roots were pruned, completely submerged plants grew less and had lower stem and leaf chlorophyll a, as compared with controls with intact roots. Roots exposed to the lowest PAR (daily mean 4·7 ± 2·4 µmol m⁻² s⁻¹) under water contained less chlorophyll, but there was no difference in aquatic root biomass after 4 weeks, regardless of light availability in the water column (high PAR was available to all emergent shoots).

Conclusions

Both M. brownii and C. coronopifolia responded to submergence with growth of aquatic adventitious roots, which essentially replaced the existing sediment root system. These aquatic roots contained chlorophyll and were photosynthetically active. Removal of aquatic roots had negative effects on plant growth during partial and complete submergence.     

Genotypic differences in Al resistance and the role of cell-wall pectin in Al exclusion from the root apex in Fagopyrum tataricum

Background and Aims

Aluminium (Al) toxicity is one of the factors limiting crop production on acid soils. However, genotypic differences exist among plant species or cultivars in response to Al toxicity. This study aims to investigate genotypic differences among eight cultivars of tatary buckwheat (Fagopyrum tataricum) for Al resistance and explore the possible mechanisms of Al resistance.

Methods

Al resistance was evaluated based on relative root elongation (root elongation with Al/root elongation without Al). Root apex Al content, pectin content and exudation of root organic acids were determined and compared.

Key Results

Genotypic differences among the eight cultivars were correlated with exclusion of Al from the root apex. However, there was a lack of correlation between Al exclusion and Al-induced oxalate secretion. Interestingly, cell-wall pectin content of the root apex was generally lower in Al-resistant cultivars than in Al-sensitive cultivars. Although we were unable to establish a significant correlation between Al exclusion and pectin content among the eight cultivars, a strong correlation could be established among six cultivars, in which the pectin content in the most Al-resistant cultivar ‘Chuan’ was significantly lower than that in the most Al-sensitive cultivar ‘Liuku2’. Furthermore, root apex cell-wall pectin methylesterase activity (PME) was similar in ‘Chuan’ and ‘Liuku2’ in the absence of Al, but Al treatment resulted in increased PME activity in ‘Liuku2’ compared with ‘Chuan’. Immunolocalization of pectins also showed that the two cultivars had similar amounts of either low-methyl-ester pectins or high-methyl-ester pectins in the absence of Al, but Al treatment resulted in a more significant increase of low-methyl-ester pectins and decrease of high-methyl-ester pectins in ‘Liuku2’.

Conclusions

Cell-wall pectin content may contribute, at least in part, to differential Al resistance among tatary buckwheat cultivars.     

Optimizing nitrogen economy under drought: increased leaf nitrogen is an acclimation to water stress in willow (Salix spp.)

Background and Aims

The major objective was to identify plant traits functionally important for optimization of shoot growth and nitrogen (N) economy under drought. Although increased leaf N content (area basis) has been observed in dry environments and theory predicts increased leaf N to be an acclimation to drought, experimental evidence for the prediction is rare.

Methods

A pedigree of 200 full-sibling hybrid willows was pot-grown in a glasshouse in three replicate blocks and exposed to two water regimes for 3 weeks. Drought conditions were simulated as repeated periods of water shortage. The total leaf mass and area, leaf area efficiency (shoot growth per unit leaf area, E_A), area-based leaf N content (N_A), total leaf N pool (N_L) and leaf N efficiency (shoot growth per unit leaf N, E_N) were assessed.

Key Results

In the water-stress treatment, shoot biomass growth was N limited in the genotypes with low N_L, but increasingly limited by other factors in the genotypes with greatest N_L. The N_A was increased by drought, and drought-induced shift in N_A varied between genotypes (significant G × E). Judged from the E_A–N_A relationship, optimal N_A was 16 % higher in the water-stress compared with the well-watered treatment. Biomass allocation to leaves and shoots varied between treatments, but the treatment response of the leaf : shoot ratio was similar across all genotypes.

Conclusions

It is concluded that N-uptake efficiency and leaf N efficiency are important traits to improve growth under drought. Increased leaf N content (area basis) is an acclimation to optimize N economy under drought. The leaf N content is an interesting trait for breeding of willow bioenergy crops in a climate change future. In contrast, leaf biomass allocation is a less interesting breeding target to improve yield under drought.     

Differences in flooding tolerance between species from two wetland habitats with contrasting hydrology: implications for vegetation development in future floodwater retention areas

Background and Aims

Plants need different survival strategies in habitats differing in hydrological regimes. This probably has consequences for vegetation development when former floodplain areas that are currently confronted with soil flooding only, will be reconnected to the highly dynamical river bed. Such changes in river management are increasingly important, especially at locations where increased water retention can prevent flooding events in developed areas. It is therefore crucial to determine the responses of plant species from relatively low-dynamic wetlands to complete submergence, and to compare these with those of species from river forelands, in order to find out what the effects of such landscape-scale changes on vegetation would be.

Methods

To compare the species'' tolerance to complete submergence and their acclimation patterns, a greenhouse experiment was designed with a selection of 19 species from two contrasting sites: permanently wet meadows in a former river foreland, and frequently submerged grasslands in a current river foreland. The plants were treated with short (3 weeks) and long (6 weeks) periods of complete submergence, to evaluate if survival, morphological responses, and changes in biomass differed between species of the two habitats.

Key Results

All tested species inhabiting river forelands were classified as tolerant to complete submergence, whereas species from wet meadows showed either relatively intolerant, intermediate or tolerant responses. Species from floodplains showed in all treatments stronger shoot elongation, as well as higher production of biomass of leaves, stems, fine roots and taproots, compared with meadow species.

Conclusions

There is a strong need for the creation of temporary water retention basins during high levels of river discharge. However, based on the data presented, it is concluded that such reconnection of former wetlands (currently serving as meadows) to the main river bed will strongly influence plant species composition and abundance.Key words: Acclimation, biomass allocation, climate, complete submergence, flooding tolerance, retention areas, shoot elongation, soil flooding, waterlogging, wetland species     

Water deficit and induction of summer dormancy in perennial Mediterranean grasses

Background and Aims

Summer dormancy is a trait conferring superior drought survival in Mediterranean perennial grasses. As the respective roles of environmental factors and water deficit on induction of summer dormancy are unclear, the effect of intense drought were tested under contrasting day lengths in a range of forage and native grasses.

Methods

Plants of Poa bulbosa, Dactylis glomerata ‘Kasbah’ and Lolium arundinaceum ‘Flecha’ were grown in pots (a) from winter to summer in a glasshouse and subjected to either an early or a late-spring drought period followed by a summer water deficit and (b) in controlled conditions, with long days (LD, 16 h) or short days (SD, 9 h) and either full irrigation or water deficit followed by rehydration. Leaf elongation, senescence of aerial tissues and dehydration of basal tissues were measured to assess dormancy. Endogenous abscisic acid (ABA) in basal tissues was determined by monoclonal immunoassay analysis.

Key Results

Even under irrigation, cessation of leaf elongation, senescence of lamina and relative dehydration of basal tissues were triggered only by a day length longer than 13 h 30 min (late spring and LD) in plants of Poa bulbosa and Dactylis glomerata ‘Kasbah’ which exhibit complete dormancy. Plants of Lolium arundinaceum ‘Flecha’ maintained leaf growth under irrigation irrespective of the day length since its dormancy is incomplete. ABA concentrations were not higher during late-spring drought than early, and could not be associated with spring dormancy induction. In summer, ABA concentration in bulbs of the desiccation-tolerant Poa were greater than in basal tissues of other species.

Conclusions

The results of both experiments tend to invalidate the hypothesis that water deficit has a role in early summer-dormancy induction in the range of tested grasses. However, a late-spring drought tends to increase plant senescence and ABA accumulation in basal tissues of forage grasses which could enhance summer drought survival.Key words: Poa bulbosa, Dactylis glomerata, Lolium arundinaceum, abscisic acid, water deficit, photoperiod, summer dormancy, perennial grass     

Characterization of a rice variety with high hydraulic conductance and identification of the chromosome region responsible using chromosome segment substitution lines

Background and Aims

The rate of photosynthesis in paddy rice often decreases at noon on sunny days because of water stress, even under submerged conditions. Maintenance of higher rates of photosynthesis during the day might improve both yield and dry matter production in paddy rice. A high-yielding indica variety, ‘Habataki’, maintains a high rate of leaf photosynthesis during the daytime because of the higher hydraulic conductance from roots to leaves than in the standard japonica variety ‘Sasanishiki’. This research was conducted to characterize the trait responsible for the higher hydraulic conductance in ‘Habataki’ and identified a chromosome region for the high hydraulic conductance.

Methods

Hydraulic conductance to passive water transport and to osmotic water transport was determined for plants under intense transpiration and for plants without transpiration, respectively. The varietal difference in hydraulic conductance was examined with respect to root surface area and hydraulic conductivity (hydraulic conductance per root surface area, L_p). To identify the chromosome region responsible for higher hydraulic conductance, chromosome segment substitution lines (CSSLs) derived from a cross between ‘Sasanishiki’ and ‘Habataki’ were used.

Key Results

The significantly higher hydraulic conductance resulted from the larger root surface area not from L_p in ‘Habataki’. A chromosome region associated with the elevated hydraulic conductance was detected between RM3916 and RM2431 on the long arm of chromosome 4. The CSSL, in which this region was substituted with the ‘Habataki’ chromosome segment in the ‘Sasanishiki’ background, had a larger root mass than ‘Sasanishiki’.

Conclusions

The trait for increasing plant hydraulic conductance and, therefore, maintaining the higher rate of leaf photosynthesis under the conditions of intense transpiration in ‘Habataki’ was identified, and it was estimated that there is at least one chromosome region for the trait located on chromosome 4.     

Relationships between xylem vessel characteristics, calculated axial hydraulic conductance and size-controlling capacity of peach rootstocks

Background and Aims

Previous studies indicate that the size-controlling capacity of peach rootstocks is associated with reductions of scion water potential during mid-day that are caused by the reduced hydraulic conductance of the rootstock. Thus, shoot growth appears to be reduced by decreases in stem water potential. The aim of this study was to investigate the mechanism of reduced hydraulic conductance in size-controlling peach rootstocks.

Methods

Anatomical measurements (diameter and frequency) of xylem vessels were determined in shoots, trunks and roots of three contrasting peach rootstocks grown as trees, each with different size-controlling characteristics: ‘Nemaguard’ (vigorous), ‘P30-135’ (intermediate vigour) and ‘K146-43’ (substantially dwarfing). Based on anatomical measurements, the theoretical axial xylem conductance of each tissue type and rootstock genotype was calculated via the Poiseuille–Hagen law.

Key Results

Larger vessel dimensions were found in the vigorous rootstock (‘Nemaguard’) than in the most dwarfing one (‘K146-43’) whereas vessels of ‘P30-135’ had intermediate dimensions. The density of vessels per xylem area in ‘Nemaguard’ was also less than in ‘P30-135’and ‘K146-43’. These characteristics resulted in different estimated hydraulic conductance among rootstocks: ‘Nemaguard’ had higher theoretical values followed by ‘P30-135’ and ‘K146-43’.

Conclusions

These data indicate that phenotypic differences in xylem anatomical characteristics of rootstock genotypes appear to influence hydraulic conductance capacity directly, and therefore may be the main determinant of dwarfing in these peach rootstocks.Key words: Prunus, rootstock, vessel diameter, hydraulic conductance, dwarfing, xylem anatomy, Poiseuille–Hagen     

The infection processes of Sclerotinia sclerotiorum in cotyledon tissue of a resistant and a susceptible genotype of Brassica napus

Background and Aims

Sclerotinia sclerotiorum can attack >400 plant species worldwide. Very few studies have investigated host–pathogen interactions at the plant surface and cellular level in resistant genotypes of oilseed rape/canola (Brassica napus).

Methods

Infection processes of S. sclerotiorum were examined on two B. napus genotypes, one resistant cultivar ‘Charlton’ and one susceptible ‘RQ001-02M2’ by light and scanning electron microscopy from 2 h to 8 d post-inoculation (dpi).

Key Results

The resistant ‘Charlton’ impeded fungal growth at 1, 2 and 3 dpi, suppressed formation of appresoria and infection cushions, caused extrusion of protoplast from hyphal cells and produced a hypersensitive reaction. At 8 dpi, whilst in ‘Charlton’ pathogen invasion was mainly confined to the upper epidermis, in the susceptible ‘RQ001-02M2’, colonization up to the spongy mesophyll cells was evident. Calcium oxalate crystals were found in the upper epidermis and in palisade cells in susceptible ‘RQ001-02M2’ at 6 dpi, and throughout leaf tissues at 8 dpi. In resistant ‘Charlton’, crystals were not observed at 6 dpi, whereas at 8 dpi they were mainly confined to the upper epidermis. Starch deposits were also more prevalent in ‘RQ001-02M2’.

Conclusions

This study demonstrates for the first time at the cellular level that resistance to S. sclerotiorum in B. napus is a result of retardation of pathogen development, both on the plant surface and within host tissues. The resistance mechanisms identified in this study will be useful for engineering disease-resistant genotypes and for developing markers for screening for resistance against this pathogen.     

Arbuscular mycorrhizal fungi alter plant allometry and biomass-density relationships

Background and Aims

Plant biomass–density relationships during self-thinning are determined mainly by allometry. Both allometry and biomass–density relationship have been shown to vary with abiotic conditions, but the effects of biotic interactions have not been investigated. Arbuscular mycorrhizal fungi (AMF) can promote plant growth and affect plant form. Here experiments were carried out to test whether AMF affect plant allometry and the self-thinning trajectory.

Methods

Two experiments were conducted on Medicago sativa L., a leguminous species known to be highly dependent on mycorrhiza. Two mycorrhizal levels were obtained by applying benomyl (low AMF) or not (high AMF). Experiment 1 investigated the effects of AMF on plant growth in the absence of competition. Experiment 2 was a factorial design with two mycorrhizal levels and two plant densities (6000 and 17 500 seeds m⁻²). Shoot biomass, root biomass and canopy radius were measured 30, 60, 90 and 120 d after sowing. The allometric relationships among these aspects of size were estimated by standardized major axis regression on log-transformed data.

Key Results

Shoot biomass in the absence of competition was lower under low AMF treatment. In self-thinning populations, the slope of the log (mean shoot biomass) vs. log density relationship was significantly steeper for the high AMF treatment (slope = –1·480) than for the low AMF treatment (–1·133). The canopy radius–biomass allometric exponents were not significantly affected by AMF level, but the root–shoot allometric exponent was higher in the low AMF treatment. With a high level of AMF, the biomass–density exponent can be predicted from the above-ground allometric model of self-thinning, while this was not the case when AMF were reduced by fungicide.

Conclusions

AMF affected the importance of below-ground relative to above-ground interactions and changed root vs. shoot allocation. This changed allometric allocation of biomass and altered the self-thinning trajectory.