The expansion of maize root-cap mucilage during hydration. 1. Kinetics

The conventional view of root-cap mucilage as an expanded blob of mucilage is characteristic only of root tips in contact with free water. In soil, the mucilage is almost always a dry coating over the tip to which soil particles adhere. The kinetics of expansion of root-cap mucilage of Zea mays roots grown in field soil, in soil in pots, and axenically on agar, were determined when the mucilage was exposed to water. On the soil-grown roots the increase in mucilage volume was linear with time, sometimes reaching a constant volume during the 6 h of measurement, but sometimes not. This linear expansion is interpreted as limited by the rate at which the condensed mucilage in the periplasmic and intercellular spaces of the root cap passes to the exterior of the cap, expanding as fast as it arrives outside in the water. The height of the plateau is interpreted as a measure of the amount of mucilage initially present in the interior spaces. Because of the greater availability of water in the axenic roots grown on 1% agar, the mucilage was already outside the root cap, and it expanded more rapidly. It reached a final volume about 10-fold greater than that on the soil-grown roots. The volume increase was curvilinear with time. An analysis of these curves suggested that this swelling on axenic roots was a diffusion of mucilage outwards from the flanks of the root cap, and the diffusivity of the mucilage was estimated as 4 × 10^?8 cm² s^?1. The molecular radius derived from this diffusivity was 34 nm, and the estimated molecular weight was 1.6 × 10⁸ Da.     

Plant and bacterial mucilages of the maize rhizosphere: Comparison of their soil binding properties and histochemistry in a model system

Mucilages from the root tips of axenically-grown maize and from a bacterium (Cytophaga sp.) isolated from the rhizosheaths of field-grown roots, were immobilized by drying onto nylon blotting membrane. The mucilage plaques remained in place through repeated rewettings and histochemical treatments. Staining of the plaques showed that both mucilages included acidic groups, and 1,2 diols (the latter notably fewer in bacterial mucilage). Bacterial mucilage plaques stained strongly for protein, plant mucilage was unstained. Plaques of both mucilages bound soil particles strongly if soil was applied to wet mucilage and then dried. Bound soil was not lost with rewetting. Dry weight and densitometer measurements showed that bacterial mucilage bound about 10% more soil than the same surface area of root-cap mucilage. Pretreatment of plaques with periodate oxidation eliminated most soil binding by root-cap mucilage but this was completely reversible by reduction with borohydride. Soil binding to bacterial mucilage was unaffected by periodate but much diminished by borohydride pretreatment (partially restored by subsequent oxidation). Neither pretreatment with cationic dyes nor preincubation in pectinase, pectin methylesterase or protease affected subsequent soil binding by the mucilage plaques. Pretreatment of root-cap mucilage plaques with lectins specific for component sugars also did not alter soil binding. It is concluded that mucilages of both plant and bacterial origin can contribute to the adhesion and cohesion of maize rhizosheaths, but each by a different mechanism. Binding by root-cap mucilage depends on 1,2 diol groups of component sugars, that of bacterial mucilage does not, and is likely to be protein mediated. ei]Section editor: R O D Dixon     

The expansion of maize root-cap mucilage during hydration. 3. Changes in water potential and water content

Root-cap mucilage from aerial nodal roots of maize has been found to have water potential values of −11 MPa or lower when air dried. The value approaches 0 MPa within 2 min of hydration in distilled water. In this time the expanding gel absorbs only about 0.3% of the water content of fully expanded mucilage. It is concluded that the root-cap mucilage per se has almost no capacity to retain water in the rhizosphere. Any function that it may play in the slowing of root desiccation would be indirect. For example, mucilage might decrease pore size between and within soil aggregates by pulling the particles together in a cycle of nocturnal efflux of water from the root surface, and diumal dyring during transpiration.     

Drying of mucilage causes water repellency in the rhizosphere of maize: measurements and modelling

Background and Aims

Although maize roots have been extensively studied, there is limited information on the effect of root exudates on the hydraulic properties of maize rhizosphere. Recent experiments suggested that the mucilaginous fraction of root exudates may cause water repellency of the rhizosphere. Our objectives were: 1) to investigate whether maize rhizosphere turns hydrophobic after drying and subsequent rewetting; 2) to test whether maize mucilage is hydrophobic; and 3) to find a quantitative relation between rhizosphere rewetting, particle size, soil matric potential and mucilage concentration.

Methods

Maize plants were grown in aluminum containers filled with a sandy soil. When the plants were 3-weeks-old, the soil was let dry and then it was irrigated. The soil water content during irrigation was imaged using neutron radiography. In a parallel experiment, ten maize plants were grown in sandy soil for 5 weeks. Mucilage was collected from young brace roots growing above the soil. Mucilage was placed on glass slides and let dry. The contact angle was measured with the sessile drop method for varying mucilage concentration. Additionally, capillary rise experiments were performed in soils of varying particle size mixed with maize mucilage. We then used a pore-network model in which mucilage was randomly distributed in a cubic lattice. The general idea was that rewetting of a pore is impeded when the concentration of mucilage on the pore surface (g cm^?2) is higher than a given threshold value. The threshold value depended on soil matric potential, pore radius and contract angle. Then, we randomly distributed mucilage in the pore network and we calculated the percolation of water across a cubic lattice for varying soil particle size, mucilage concentration and matric potential.

Results

Our results showed that: 1) the rhizosphere of maize stayed temporarily dry after irrigation; 2) mucilage became water repellent after drying. Mucilage contact angle increased with mucilage surface concentration (gram of dry mucilage per surface area); 3) Water could easily cross the rhizosphere when the mucilage concentration was below a given threshold. In contrast, above a critical mucilage concentration water could not flow through the rhizosphere. The critical mucilage concentration decreased with increasing particle size and decreasing matric potential.

Conclusions

These results show the importance of mucilage exudation for the water fluxes across the root-soil interface. Our percolation model predicts at what mucilage concentration the rhizosphere turns hydrophobic depending on soil texture and matric potential. Further studies are needed to extend these results to varying soil conditions and to upscale them to the entire root system.

     

The expansion of maize root-cap mucilage during hydration. 2. Observations on soil-grown roots by cryo-scanning electron microscopy

Expansion of root-cap mucilage during hydration was followed by cryo-scanning analytical microscopy of soil-grown roots of diploperennis and Zea mays. Roots examined directly from the soil have no expanded mucilage. Their condensed, unexpanded mucilage is in three domains, periplasmic, intercellular and peripheral to the cap tissue. Carbon concentration is the same in the three domains. During hydration there is no change in carbon concentration as the condensed mucilage moves through these three domains; however there is a sharp drop at the periphery where a gel phase transition occurs. The rate of expansion of the mucilage blob around the root tip is limited by the rate of this gel phase transition.     

Do detached root-cap cells influence bacteria associated with maize roots?

Abstract. A model rhizosphere has been used which consisted of detached root-cap cells of maize in their surrounding root-cap mucilage on the surface of Noble agar. These cells were co-cultured for periods up to 32 d with eight different bacterial isolates from soil-grown roots and surrounding soil and two laboratory cultures. Cap cells were unaffected by the bacteria. There were five different type-specific responses of the bacteria in proximity to the cap cells. There were, strong growth inhibition ( Rhizobium sp. and Escherichia coli ), strong stimulation ( Pseudomonas fluorescens , laboratory strain), mixed weak inhibition or stimulation ( Pseudomonas fluorescens , field isolate), early inhibition followed by strong stimulation then spore formation ( Bacillus spp.), no effect ( Streptomyces sp. and Cytophaga sp.). It is concluded that detached root-cap cells are actively involved in the establishment of characteristic rhizosphere bacterial microflora.     

朱唇种子吸水特性及其在干旱胁迫下萌发特性

种子粘液质是植物在长期适应环境过程中形成的,该物质对于种子的扩散、定居、生存力的改善、萌发、幼苗生存乃至抵御有毒化学物质毒害等都具有重要的生态学意义。朱唇为唇形科鼠尾草属多年生草本植物,原产美洲热带地区,现已广泛栽植于世界各地。为了理解朱唇种子表面的粘液物质吸水特性和种子在干旱胁迫下的萌发特性,该研究以朱唇种子为材料,运用光学显微镜和扫描电子显微镜观察以及种子萌发试验的方法,对种子和粘液层的形态结构、粘液质对种子萌发的影响进行了研究。结果表明:朱唇种子为卵形,表面为负网状结构,千粒重为(1.611±0.0084)g,无粘液种子吸水倍数为3,粘液种子吸水倍数为25,粘液层吸水倍数为122。粘液和无粘液种子及粘液层的重量都随吸水时间的延长而增长,但脱水过程要远长于吸水过程。朱唇种子吸水2 h达到饱和,经过36 h可干燥失水恢复原重。不同浓度PEG对朱唇种子的萌发均有影响,发芽势随PEG浓度升高而显著降低。朱唇种子在5%PEG胁迫下发芽率最高达(90.00±8.66)%,20%PEG胁迫下发芽率最低为(76.67±10.41)%,低浓度PEG对朱唇种子萌发有一定促进作用。这说明朱唇种子为速萌型种子,其粘液质在种子吸水过程中起到举足轻重的作用,能保证短时间内有充足的水分供其萌发。     

A new approach for the quantification of root-cap mucilage exudation in the soil

Direct evidence on the functions of root-cap mucilage during plant root growth in soil is limited mainly due to the lack of a method for in situ measurements. In this paper, we offer a method that facilitates the measurement of mucilage exudation when roots are growing in soil. We observed the mucilage exudation directly through a transparent panel located on the side of a root box in which plant roots were growing. We used a CCD camera attached to a microscope to observe and record mucilage exudation. Using image analysis, the activity of mucilage exudation was evaluated based on the area occupied by the mucilage on the root tip. The area of mucilage observed on the root tips after 1-h growth in soil corresponded with the weight of mucilage that was originally observed on the tips before they were transplanted. This relationship suggests that the observed area on root tip relates to total exudation. The area of mucilage exudation on the root tips was high (0.48 mm²) at night and low (0.35 mm²) at midday, suggesting that the activity of mucilage exudation follows diurnal changes. Furthermore, the mucilage exudation positively correlated with the root elongation rate, implying that fast-growing roots exude more mucilage.     

The role of seed provenance in the early development of Arbutus unedo seedlings under contrasting watering conditions

In the last decades, several studies have reported the increase of land degradation and desertification in the Mediterranean Basin. Depending on degradation severity, ecological restoration might be needed in order to promote ecosystem recovery. The ecology of the selected species and intra specific variability should be considered in order to improve restoration options, especially facing climate change.The present study tested the hypothesis that seedlings from drier provenances would be better adapted to low water content conditions. Seeds were germinated under controlled temperature after which seedlings were grown in a phytotron under two contrasting watering regimes. Seedling performance was analysed using morphological and physiological parameters.Low water content had a clear negative effect on the seedlings’ aboveground biomass (total dry weight, root collar diameter, leaf dry weight and leaf weight ratio) and a positive effect on belowground biomass (root weight and root:shoot ratio). This response was not unequivocal, since provenances differed in morphological adaptations to low water content. Seedlings from the wettest provenance revealed a higher relative growth rate under high water content but a poor adaptation to limited water availability when compared to the other two provenances. This was observed by the absence of a significant belowground investment in this provenance. Seedlings from the wettest provenance also presented a significant reduction of total leaf area that was not observed in the other two provenances. This can however be hardly considered as a successful adaptation to cope with drought since this provenance produced less sclerophyllous leaves, less belowground biomass and also lower sapwood to leaf area ratio independently from the water content conditions. By contrast, seedlings from the dry provenance with the hottest summer had similar root collar diameter, leaf dry weight and physiological performance under both watering regimes.The observed adaptations to water regimes seem to be related with the climate of the seed source and highlighted the importance of seed provenance in ecological restoration actions using Mediterranean species. This knowledge could improve early establishment success predictions for different plant populations, allowing more reliable and cost-effective management decisions under climate change scenarios.     

Effects of evening and nighttime leaf wetting on stomatal behavior of Cryptomeria japonica growing in dry soil

To examine the hypothesis that stomatal behavior of plants in dry soil is influenced by a slow recovery from daytime water deficit, we studied the effect of repeated wetting of leaves during evening and night in Cryptomeria japonica seedlings grown in dry soil. After 7 and 10 days of leaf wetting treatment the midday leaf water potential decreased and the transpiration rate increased, respectively. Therefore, we suggest that rapid recovery from daytime water deficit could weaken the water conserving stomatal behavior that adapts to drought conditions in the roots.     

小麦幼苗根系形态与反复干旱存活率的关系

以35个不同栽培类型的小麦品种（系）作为试验材料,根据其6叶幼苗的根系形态性状进行聚类分析,供试材料的根系类型分为3种：大根系、小根系和中间型根系。具有中间型根系的材料反复干旱存活率最高,这些材料的根系特点是单株根数7－8．5条,最大根长20-22cm,根总干重44－48mg,其中10cm以下根干重占36％－45％,根冠比范围在0.22-0.24。一些水地栽培的育成品种苗期抗旱性较强,旱地栽培的育成品种苗期抗旱性差异较大,个别旱地栽培的地方品种在土壤水分胁迫条件下反而比在正常水分条件下的根系发育更好,可能是长期适用干旱条件的结果。     

新疆短命植物抱茎独行菜种子粘液质特性的研究

以新疆荒漠植物抱茎独行菜为材料,运用光镜与扫描电镜观察以及紫外吸收光谱法、化学反应及种子萌发实验等方法,对粘液质的形态和结构,物理化学特性,粘液质对种子萌发及萌发后的影响进行了研究.结果显示:(1)完整干种子表面覆盖着一层膜状物质(完全脱水的粘液质),并呈同一走向的山脊状突出的网状结构,遇水后粘液物质呈射线状向外发射出来,化学反应实验结果表明,粘液质的组成可能是某种多糖,如β-葡聚糖.(2)粘液质约占干种子重量的1/4,有很强的吸水能力,完全浸润10 min后,种子重量增加约30~40倍,种子长度、宽度、厚度的增加分别多于1倍、2倍、4倍;完全润湿的种子能够粘附相当于其干种子重量68倍的沙粒.(3)种皮粘液质对于不同土壤基质中的种子萌发有重要作用,但是对萌发后幼苗的生长没有作用.     

Exogenous salicylic acid alleviates water stress in watermelon plants

Salicylic acid (SA) has been considered to attenuate the effects of abiotic stresses on plants, including water deficit that intensely affects the growth and production of plants. The goal of this work was to evaluate the role of SA in the alleviation of water stress in watermelon seedlings on a morphophysiological and biochemical level. The experiment consisted of application of SA at concentrations of 0, 0.25, 0.50, 0.75 and 1.0 μmol L⁻¹ to leaves of watermelon seedlings grown in three levels of water retention (100%, 75% and 50% WRL). To evaluate the effect on morphophysiological and biochemical aspects, the plant height, leaf area, shoot and root dry weight, chlorophyll index, relative water content, electrolyte leakage, protein content, amino acids, proline, carbohydrates, sucrose and starch concentration variables were determined. All variables were influenced by the SA concentrations and WRL, with statistically significant interaction between these factors for all except root dry weight. SA promotes increases in the concentration of organic solutes and reduces the rate of electrolyte leakage in watermelon seedlings, thus, supporting metabolism and growth of plants under stress conditions resulting from water deficit.     

高产杂交稻两优培九旱育秧苗期抗氧化系统活性的研究

以汕优63为对照,研究了旱育条件下两优培九幼苗的体内抗氧化系统的特性。显示两优培九在旱育条件下其体内三大抗氧化酶SOD、POD和CAT活性都较水育条件下高,而体内超氧阴离子含量和MDA含量和水育秧相比却没有大的差异;旱育条件下秧苗叶片内积累的脯氨酸量显著高于水育秧。这不但能提高水稻苗期抗旱能力,同时也为移栽大田后具有明显的生长优势和抗胁迫能力打下了物质基础。另外,发现秧苗培育时间过长也不利于获得高质量的秧苗,时间以30 d左右为宜。和汕优63相比,相同条件下的两优培九幼苗抗氧化系统活性较高,这也可能是两优培九高产的生理基础之一。     

Mucilages and polysaccharides in Ziziphus species (Rhamnaceae): localization, composition and physiological roles during drought-stress.

The drought-tolerant tree species Ziziphus mauritiana Lamk. and Z. rotundifolia Lamk. were shown to have similar high mucilage concentrations (7-10% dry weight) in their leaves, with large numbers of mucilage-containing cells in the upper epidermis and extracellular mucilage-containing cavities in the leaf veins and stem cortex. The main sugar constituents of the water-soluble mucilage extract were rhamnose, glucose and galactose. During drought-stress in two independent studies, foliar mucilage content was unaffected in both species, but glucose and starch contents declined significantly in crude mucilage extracts from droughted leaves. Enzymatic hydrolysis of the mucilage extract using alpha-amylase and amyloglucosidase released glucose, indicating that a mucilage-associated water-soluble glucan, with alpha-1,4- and alpha-1,6-linkages, may exist which was extracted together with the mucilage. From the current data, it is not possible to localize the glucan to determine whether or not it is associated with mucilage-containing cells. Data from pressure-volume analyses of drought-stressed and control leaves showed that, in line with their similar mucilage contents, the relative leaf capacitance isotherm (change in relative water content per unit change in water potential) was similar in both species. During drought-stress, reduced relative capacitance resulted from osmotic adjustment and decreased wall elasticity. Data suggest that in Ziziphus leaves, intracellular mucilages play no part in buffering leaf water status during progressive drought. In Ziziphus species, growing in environments with erratic rainfall, the primary role of foliar mucilage and glucans, rather than as hydraulic capacitors, may be as sources for the remobilization of solutes for osmotic adjustment, thus enabling more effective water uptake and assimilate redistribution into roots and stems prior to defoliation as the drought-stress intensified.     

Spring water deficit and soil conditions matter more than seed origin and summer drought for the establishment of temperate conifers

In anticipation of more severe summer droughts, forestry in temperate Europe is searching for drought-resistant ecotypes of native tree species that might maintain ecosystem services in the future. We investigated how spring precipitation and soil conditions interact with summer drought and affect the establishment of conifer seedlings from different climatic origin. Emergence, establishment and subsequent performance of seedlings originating from autochthonous, Central Alpine, continental Eastern European, and Mediterranean Pinus sylvestris and Picea abies populations were studied in the dry Alpine Rhine valley, Switzerland, at three sites with differing soil water holding capacities and in 3 years with contrasting weather conditions. In addition to this natural inter-annual variation, precipitation was manipulated within sites with throughfall reduction roofs. Seedling establishment and growth were principally affected by the spring weather in the year of emergence. In years with average to positive spring water balance, seedlings grown at the site with the highest water holding capacity had 2–5 times more aboveground biomass than seedlings grown at sites with less favourable soils. Effects of seed origin were marginal and only detectable at the drier sites: contrary to our expectations, seedlings from the Central Alpine Rhone valley, where the climatic spring water deficit is large, outperformed those from the Mediterranean. Consequently, plantation of non-native populations from dryer origin will mitigate the effects of increased summer drought at driest sites only, while the inter-annual variability of spring precipitation will continue to enable temperate conifers to regenerate on a wide range of forest soils independent of seed origin.     

Acclimatizing micropropagated black cherry by comparison with half-sib seedlings

Black cherry ( Prunus serotina Ehrh.) plantlets were cultured in vitro, transferred to potting mixture and evaluated for 8 weeks for acclimatization to a varying, but controlled environment. Whole plantlet growth and water relations were monitored and compared to seedlings of comparable size of the same maternal genotype grown under different conditions, but given the same pretest environment. At one week ex vitro. gravimetrically determined leaf conductance of plantlets was high, but became closer to that of seedlings in both magnitude and diurnal pattern as acclimatization progressed. By 8 weeks, leaf conductance of plantlets was nearly identical to that of seedlings, but the xylem water potential of the plantlets was significantly less. Logarithmic regressions of shoot vs root dry weight indicated that seedlings were allocating twice as much dry matter to shoot than to root growth compared to 4- and 8-week plantlets. Over the same period, stomatal densities of both seedlings and plantlets decreased and stomatal pore lengths increased. Multiple adventitious roots of plantlets emerged from a single site just below the root collar whereas secondary or lateral roots of seedlings originated acropetally along the central root axis. Leaf conductance of plantlets at 8 weeks was similar to that of seedlings indicating satisfactory acclimatization. The larger relative root growth rate of plantlets compared to the shoot, however, was associated with lower stem xylem water potential. The anomaly may be a consequence of the method of in vitro root formation. Plantlet growth rates were lower than growth rates of seedlings and their leaf area was correspondingly less.     

Plasticity of rhizosphere hydraulic properties as a key for efficient utilization of scarce resources

Background

It is known that the soil near roots, the so-called rhizosphere, has physical and chemical properties different from those of the bulk soil. Rhizosphere properties are the result of several processes: root and soil shrinking/swelling during drying/wetting cycles, soil compaction by root growth, mucilage exuded by root caps, interaction of mucilage with soil particles, mucilage shrinking/swelling and mucilage biodegradation. These processes may lead to variable rhizosphere properties, i.e. the presence of air-filled gaps between soil and roots; water repellence in the rhizosphere caused by drying of mucilage around the soil particles; or water accumulation in the rhizosphere due to the high water-holding capacity of mucilage. The resulting properties are not constant in time but they change as a function of soil condition, root growth rate and mucilage age.

Scope

We consider such a variability as an expression of rhizosphere plasticity, which may be a strategy for plants to control which part of the root system will have a facilitated access to water and which roots will be disconnected from the soil, for instance by air-filled gaps or by rhizosphere hydrophobicity. To describe such a dualism, we suggest classifying rhizosphere into two categories: class A refers to a rhizosphere covered with hydrated mucilage that optimally connects roots to soil and facilitates water uptake from dry soils. Class B refers to the case of air-filled gaps and/or hydrophobic rhizosphere, which isolate roots from the soil and may limit water uptake from the soil as well water loss to the soil. The main function of roots covered by class B will be long-distance transport of water.

Outlook

This concept has implications for soil and plant water relations at the plant scale. Root water uptake in dry conditions is expected to shift to regions covered with rhizosphere class A. On the other hand, hydraulic lift may be limited in regions covered with rhizosphere class B. New experimental methods need to be developed and applied to different plant species and soil types, in order to understand whether such dualism in rhizosphere properties is an important mechanism for efficient utilization of scarce resources and drought tolerance.     

Consumption of wheat seed reserves during germination and early growth as affected by soil water potential

Seed size and weight are important criteria for determining seedling vigour and stand establishment. Evolution of seed dry weight of wheat (Triticum aestivum L.) during germination and early growth was examined because poor stands are often associated with the depletion and exhaustion of seed reserves. Two laboratory experiments were conducted on filter paper and in soil at three water potentials using wheat seeds. Seed, root, and shoot dry weights were recorded at approximately one-day intervals. Coleoptile and first leaf lengths were also measured at all sampling periods. Wheat seedlings grown on filter paper in the dark grew to a length of 90 to 100 mm with 50% of the initial seed weight remaining after eight days when the experiment was terminated. In soil, wheat seedlings grew 15 mm with 25% of the initial seed weight remaining. Seed reserves were depleted more quickly when the soil was wet because seedlings grew more quickly. There were significant and similar negative relationships between seed weight and coleoptile length of wheat seedlings grown on filter paper and in soil. There was no effect of soil water potential on the relationship between seed weight and shoot length. Therefore, it was concluded that poor wheat stands are not likely to occur due to depletion of seed reserves under field conditions without mechanical obstacles.     

盐旱互作对不同生境盐地碱蓬种子萌发和幼苗生长的影响

研究了盐旱互作对潮间带和盐碱地生境盐地碱蓬棕色种子萌发、地上部生长和离子积累的影响。不同盐浓度预处理后,未萌发的种子风干后复水,其萌发率与对照相比没有降低,说明两种生境盐地碱蓬种子萌发期间都耐干湿交替。400mmol/LNaCl溶液浇灌的潮间带生境盐地碱蓬幼苗在第3次干旱处理后萎蔫幼苗的百分比高于盐碱地生境的,而复水后正常幼苗的百分比却相反。400mmol/LNaCl溶液处理下,第3次干旱处理复水后,盐碱地生境盐地碱蓬幼苗地上部离子含量(主要是Na+和Cl-)高于潮间带生境的。表明在干旱情况下,盐碱地生境盐地碱蓬幼苗能积累更多的无机离子,降低渗透势,提高根系吸收水分的能力。上述结果说明,盐碱地生境盐地碱蓬幼苗较潮间带生境盐地碱蓬幼苗更耐盐与干旱的交互作用。