Scanning electron microscopic aspects of short tuberized roots, with special reference to cell rhizodermis evolution under drought and rehydration

Abstract. Different morphological aspects of short tuberized roots initiated during drought stress in mesophytic species, such as Sinapis alba L., were studied with the scanning electron microscopy technique of cryofixation. Specially adapted for direct and immediate observation of fresh living roots, this method has given precise information about rhizodermal organization and evolution during drought and rehydration.
The main difference from a normal lateral root grown in a well-watered soil appears in the basal enlarged zone of the short root where all the cells show the same round and turgid aspect. In the medium zone of the short root, rhizodermis differentiation into alternating rows of short (trichoblasts) and long (atrichoblasts) cells, which characterizes the typical Sinapis root, remains clearly discernible, though not so regular as in normal lateral roots.
The turgid state of rhizodermal cells all over the short tuberized root grown in a drying soil suggests an effective regulatory mechanism for water deficit avoidance.
During the first hours of rehydration, immediate absorption of water is noticeable through the rapid swelling of some long cells which appear to protrude considerably among other rhizodermal cells. However, these protrusions will not give rise to hairs, as further observations of short roots after growth has resumed show very distinctly that progressive hair formation occurs in the medium zone of the root, and that their emergence originates from trichoblasts only.
These observations may indicate that atrichoblasts, on account of their highly vacuolated condition, are the first cells to absorb water and that they may even be stimulated, in some environmental conditions, to initiate hair formation, although they are not so well adapted to do so as the short cells in this species.     

Partial floral evocation by high irradiance in the long-day plant Sinapis alba

Previous work has shown that increased irradiance promotes flower initiation in the long-day plant Sinapis alba when given during the first half of a single, suboptimal long day. It is without flower-promoting effect, however, when given during a single short day. In the present work it is shown that, in the apex of plants exposed to a single short day at high irradiance, some changes occur that are normally observed during the transition to flowering (full evocation), e.g., elevated soluble sugar and starch levels, increased numbers of mitochondria and changed nucleolus structure. These changes are of similar magnitude and follow the same sequence as the corresponding changes during full evocation. Other changes, normally associated with full evocation, e.g., increased mitotic activity, are not caused by one short day at high irradiance. This treatment thus produces only what we call "partial evocation".     

Drought effects on polyphenol composition and antioxidant activities in aerial parts of <Emphasis Type="Italic">Salvia officinalis</Emphasis> L.

Salvia officinalis L. is a medicinal plant containing several compounds with important pharmacological activity. In this study, we investigated the effects of water deficit (moderate and severe water deficits) on the contents of total and individual polyphenols of the aerial parts. Also, we studied the effect of drought on the antioxidant activity of methanolic extracts. Our results showed that water deficiency, as estimated by the decrease in water potential, resulted in a reduction of the biomass, plant height and total chlorophyll contents. In general, drought increased the level of total and individual polyphenols and this increase was more pronounced under moderate water deficit. These findings suggest that S. Officinalis is a sensitive species and that a severe water deficit could result in a decline in the activity of enzymes involved in the biosynthesis of phenolic compounds. On the other hand, our results showed an enhancement of reducing power and the radical scavenging activity as assessed using the DPPH assay with increasing stress severity. Finally, the evaluation of the chelating capacity of the extracts was found to be altered significantly under severe treatment by 39.71%. Based on these results, it seems that drought tolerance of S. officinalis is related to the capacity of the plant to modulate its phenolics in order to face to oxidative stress caused by water limiting conditions.     

Drought adaptability of Agrobacterium rhizogenes-induced roots in oilseed rape (Brassica napus var. oleifera)

Abstract. Soil grown oilseed rape ( Brassica napus L. var. oleifera M., cv. Darmor) seedlings at the cotyledon stage (one week old), were inoculated in vivo at the base of the hypocotyl with Agrobacterium rhizogenes harbouring the pRi 15834 plasmid. Resulting adventitious root formation was observable about 2 or 3 weeks after infection. Differential Ri-induced root emergence and subsequent development occurred depending on water conditions and closeness of the wounding site to the soil surface: either thin, hairy roots growing rapidly and plagiotropically at the soil level under humid atmosphere, or hairless and fleshy, slowly growing aerial roots developed. The hairy roots were highly drought susceptible, whereas aerial roots revealed some potential for drought tolerance. Unlike normal roots, none of these Ri-induced roots appeared able to give rise to drought rhizogenesis in plants subjected to progressive drought stress. However, under hardening, achieved through successive and moderate drought stress-rehydration cycles, both types of Ri-induced roots improved drought tolerance and could express the morphogenetic differentiation programme leading to the formation of short, tuberized, drought-adapted, roots. These results, discussed in terms of hormonal imbalance and drought tolerance regulation, suggest that the Ri T-DNA gene expression, responsible for adventitious root induction and growth behaviour, is further regulated through the host plant.     

Differences in growth and water relations among Phaseolus vulgaris cultivars in response to induced drought stress

Relatively little ecophysiological research has been conducted to determine the responses to drought of Phaseolus vulgaris. Four bean cultivars (cvs.) from Brazil, A320, Carioca, Ouro Negro and Xodó were submitted to an imposed water deficit in order to evaluate the importance of some adaptive mechanisms of drought resistance through the analysis of growth parameters, water status, gas exchange and indicators of tolerance mechanisms at the cellular level. During the drought treatment, relative growth rates were more reduced for A320 and Xodó than Carioca and Ouro Negro. A320 closed its stomata very rapidly and complete stomatal closure was obtained at Psi(w)=-0.6 MPa, in contrast to the other cvs. where stomata were fully closed only at Psi(w)=-0.9 MPa. Net assimilation rates were closely related to stomatal conductances. Mechanisms at the cellular level appeared to be mostly important for higher tolerance. Carioca and Ouro Negro, when compared to A320 and Xodó, were characterized by having better drought tolerance mechanisms and higher tissue water retention capacity leading to a better growth under water deficits. The leaf dehydration rates of those cvs. were slow whereas those of the drought sensitive cvs. were rapid. The results were confirmed by the electrolyte leakage test and leaf osmotic potential measurements, which indicated higher membrane resistance and osmotic adjustment in the two tolerant cvs. Carioca and Ouro Negro. It appears from this study that despite being cultivated in the same geographical region, the four cvs. of P. vulgaris displayed somewhat different drought adaptive capacities for prolonged drought during the vegetative phase.     

Morphological and physiological responses of <Emphasis Type="Italic">Vitex negundo</Emphasis> L. var. <Emphasis Type="Italic">heterophylla</Emphasis> (Franch.) Rehd. to drought stress

Water is a main factor limiting plant growth. Integrative responses of leaf traits and whole plant growth to drought will provide implications to vegetation restoration. This study investigated the drought responses of Vitex negundo L. var. heterophylla (Franch.) Rehd. with a focus on leaf morphology and physiology, seedling growth and biomass partitioning. Potted 1-year-old seedlings were subjected to four water supply regimes [75, 55, 35 and 15% field capacity (FC)], served as control, mild water stress, moderate water stress and severe water stress. Leaf morphological traits varied to reduce the distance of water transfer under water stress and leaflets were dispersed with drought. Net photosynthetic rate decreased significantly under water stress: stomatal closure was the dominant limitation at mild and moderate drought, while metabolic impairment was dominant at severe drought. The physiological impairment at severe drought could also be detected from the relative lower water use efficiency and non-photochemical quenching to moderate water stress. Total biomass of well-watered plants was more than twice that at moderate water deficit and nearly ten times that at severe water deficit. In summary, V. negundo var. heterophylla had adaptation mechanism to water deficit even in the most serious condition, but different strategies were adopted. Seedlings invested more photosynthate to roots at mild and moderate drought while more photosynthate to leaves at severe drought. A nearly stagnant seedling growth and a sharp decline of total biomass were the survival strategy at severe water stress, which was not favorable to vegetation restoration. Water supply above 15% FC is recommended for the seedlings to vegetation restoration.     

Time-dependent changes in the responsiveness to light of phytochrome-mediated anthocyanin synthesis

Abstract. It is well established that seedlings of mustard ( Sinapis alba L.) synthesize juvenile anthocyanin only if treated with light pulses or continuous light. The light effects are considered to be due to the operation of phytochrome. Here we show that the responsiveness of anthocyanin synthesis to a saturating red light pulse or to continuous far-red light varies as a function of time and is strongly influenced by a light pretreatment prior to competence. Competence appears approximately 25 h after sowing. The starting point of anthocyanin synthesis, which is 27 h after sowing, and the lag- phase of this response, which is 2 h, are not affected by light pretreatments prior to competence. It is concluded that quantitative interpretations of phytochrome responses based entirely on properties of phytochrome can no longer be considered adequate.     

Effects of irradiation level on leaf growth of sunflower

Sunflower, Helianthus annuus L. cv. INRA 6501, plants were grown in a gravel culture subirrigated with Hoagland nutrient solution, at photosynthetically active radiation levels of 15, 30 and 60 W m^-2 at a daylength of 16 h, a temperature of 20°C and a relative humidity of 60% throughout. Development of the plant and growth of the leaves were measured. High irradiance accelerated development proportionally in all phases from germination, through leaf initiation, primordial flower formation and the maturation of all plant organs until anthesis. High irradiance levels stimulated the expansion of the growing shoot, which produced more and larger primordia. Under constant conditions the ratio between leaf initiation rate and mature length of a leaf remained constant, although the growth patterns [relationship between relative growth rate (RGR) and organ age] of successive leaves were not similar. Consequently, it may be assumed that, as in poplar, the increasing size of the growing shoot reflects the increase of the vascular system of sunflower. The growth patterns of the leaves depend on the developmental stage of the plant and, in the young primordial stage, also on irradiance level. In the linear phase of growth the growth pattern is independent of irradiance level.     

Climate-tree-growth relationships of European beech (Fagus sylvatica L.) in the French Permanent Plot Network (RENECOFOR)

The influence of climate on the radial growth of Fagus sylvatica was investigated using 15 chronologies developed from mature stands of the French Permanent Plot Network (RENECOFOR) growing under different climatic and soil conditions. The relationships between climate and ring widths were analyzed using extreme growth years, simple correlations and response functions analysis. Monthly climatic regressors were derived by a physiological water balance model that used daily climatic data and stand parameters to estimate soil water deficits. The three most frequent negative pointer years (1959, 1989, 1976) result from a particularly intense and durable drought, whereas positive years (1977, 1958) coincide with wet conditions. The total ring chronology variance attributable to climate averages 34.1% (15.8% –57%). Current early-summer soil water deficit enters in 10 models and the deficit in June explains alone a large part of the radial growth variability (mean value: 26.6%). Temperature or soil water deficit for the other months and weather conditions during the previous season were of little consistency across stands. The response pattern of earlywood is very similar and the percentage of variance explained is higher (16.2% –57.8%). Latewood widths present a different response pattern. High minimum temperature in August and/or September often favour wide latewood widths and monthly water deficits play a secondary role. The percentage of variance explained ranges from 8.8% to 67.4%. Soil water capacity strongly modulates ring characteristics and climate-growth relationships. Mean sensitivity, expressed population signal, signal-to-noise ratio and the strength of growth-climate correlations increase with decreasing soil water capacity.     

Photocontrol of the cotyledonary blade epinasty of Sinapis alba seedlings. Effects of irradiance and quality of light

The effects of the fluence rate of continuous light (far-red, fluorescent white, blue and red light) on the increase of epinastic percentage of cotyledons were recorded for six days. The epinastic photoresponse was compared to the cotyledonary enlargement and to the inhibition of the hypocotyl extension of irradiated Sinapis alba L. seedlings. Fluence rate response curves and wavelength response curves showed that the photoepinasty is a typical high irradiance response (as is the inhibition of hypocotyl extension) with two maxima, one in the blue and one in the far-red. The epinastic photoresponse was not directly related to cotyledonary growth, which was greater in red and far-red light than in blue light.     

Deficit irrigation promotes arbuscular colonization of fine roots by mycorrhizal fungi in grapevines (<Emphasis Type="Italic">Vitis vinifera</Emphasis> L.) in an arid climate

Regulated deficit irrigation (RDI) is a common practice applied in irrigated vineyards to control canopy growth and improve fruit quality, but little is known of how imposed water deficits may alter root growth and colonization by beneficial arbuscular mycorrhizal fungi (AMF). Thus, root growth and mycorrhizal colonization were determined throughout the growing season for 3 years in own-rooted, field-grown, ‘Cabernet Sauvignon’ grapevines exposed to three RDI treatments. Vines under standard RDI were irrigated at 60 to 70% of full-vine evapotranspiration (FVET) from 2 weeks after fruit set until harvest, a standard commercial practice. Early deficit vines were exposed to a more extreme deficit (30% FVET) during the period from 2 weeks after fruit set until the commencement of ripening (veraison), and thereafter reverted to standard RDI. Late deficit vines were under standard RDI until veraison, then exposed to a more extreme deficit (30% FVET) between veraison and harvest. The production of fine roots was reduced in both the early and late deficit treatments, but the reduction was more consistent in the early deficit vines because the additional deficit was imposed when roots were more rapidly growing. The frequency of arbuscules in fine roots was greater in both of the additional deficit treatments than in the standard RDI, a response that appeared chronic, as the higher frequency of arbuscules was observed throughout the season despite the additional deficits being applied at discrete times. It appears that grapevines compensated for a lower density of fine roots by stimulating arbuscular colonization. Irrigation did not affect yield or quality of grapes, but reduced whole-vine photosynthesis during the additional deficit periods. It appears that high-quality grapes can be produced in this region with less water than that applied under the current RDI practice because the root system of the vine may be more efficient due to greater arbuscular colonization by AMF. The use of trade, firm, or corporation names in this publication is for the information and convenience of the reader. Such use does not constitute an official endorsement or approval by the United States Department of Agriculture or the Agricultural Research Service of any product or service to the exclusion of others that may be suitable.     

华北地区冬小麦灌溉制度及其环境效应研究进展

充分利用有限的灌溉水资源确保冬小麦安全生产是华北地区冬小麦稳产高产面临的严峻挑战,解决这一问题的关键在于如何基于环境效应科学地进行灌溉管理。综述了国内外有关冬小麦的灌溉管理制度,即充分灌溉与非充分灌溉管理制度以及冬小麦关键灌溉期的环境效应,在此基础上提出了华北地区冬小麦科学灌溉拟重点关注的研究任务:(1)冬小麦生长发育需水时间与需水量的控制机制研究;(2)冬小麦干旱发生发展过程与致灾临界气象条件研究;(3)气候变化背景下极端干旱事件的冬小麦脆弱性诊断与适应性管理,以为华北地区冬小麦安全生产措施制定提供依据。     

Water Deficit and Spatial Pattern of Leaf Development. Variability in Responses Can Be Simulated Using a Simple Model of Leaf Development

We analyzed the effect of short-term water deficits at different periods of sunflower (Helianthus annuus L.) leaf development on the spatial and temporal patterns of tissue expansion and epidermal cell division. Six water-deficit periods were imposed with similar and constant values of soil water content, predawn leaf water potential and [ABA] in the xylem sap, and with negligible reduction of the rate of photosynthesis. Water deficit did not affect the duration of expansion and division. Regardless of their timing, deficits reduced relative expansion rate by 36% and relative cell division rate by 39% (cells blocked at the G0-G1 phase) in all positions within the leaf. However, reductions in final leaf area and cell number in a given zone of the leaf largely differed with the timing of deficit, with a maximum effect for earliest deficits. Individual cell area was only affected during the periods when division slowed down. These behaviors could be simulated in all leaf zones and for all timings by assuming that water deficit affects relative cell division rate and relative expansion rate independently, and that leaf development in each zone follows a stable three-phase pattern in which duration of each phase is stable if expressed in thermal time (C. Granier and F. Tardieu [1998b] Plant Cell Environ 21: 695–703).     

Effects of soil and air drought on growth,plant water status and leaf gas exchange in three Mediterranean cedar species: <Emphasis Type="Italic">Cedrus atlantica</Emphasis>, <Emphasis Type="Italic">C. brevifolia</Emphasis> and <Emphasis Type="Italic">C. libani</Emphasis>

Three- and four-year-old potted, greenhouse-grown cedar seedlings were subjected to two different watering regimes: half received full water supply and the other half was submitted to moderate drought (50% of the full water supply). Height growth was the greatest for C. atlantica and the most-limited for C. brevifolia in the well-watered set. However, in the dry set, height growth was less affected by drought conditions for C. brevifolia than for C. atlantica. Cedrus libani gave intermediate results for both watering regimes. Moderate drought provoked a decrease in osmotic potential at full leaf turgor and a long-lasting osmotic adjustment. When irrigation was withheld completely to induce severe soil drying, gas exchange decreased and then stopped at predawn water potentials of −3.0 MPa for C. brevifolia, between −2.6 and −2.8 MPa for C. libani, and at −2.4 MPa for C. atlantica, irrespective of watering regime. For all species, the dry set showed lower net photosynthesis (A) and stomatal conductance (g _s) than the plants in the well-watered set. A and g _s responded to variations in atmospheric water-vapour pressure deficit (VPD). As VPD increased, A and g _s decreased, and this trend was proportionate to initial values at low VPD, but remained independent of previous watering treatments, plant water status or species. To conclude, C. brevifolia appears to be a species with limited growth potential but strong soil drought tolerance whereas C. atlantica has strong growth potential when an adequate water supply is available but is more sensitive to soil drought. C. libani shows an intermediate behaviour for growth and drought tolerance.     

Water deficits uncouple growth from photosynthesis, increase C content, and modify the relationships between C and growth in sink organs

In plants, carbon (C) molecules provide building blocks for biomass production, fuel for energy, and exert signalling roles to shape development and metabolism. Accordingly, plant growth is well correlated with light interception and energy conversion through photosynthesis. Because water deficits close stomata and thus reduce C entry, it has been hypothesised that droughted plants are under C starvation and their growth under C limitation. In this review, these points are questioned by combining literature review with experimental and modelling illustrations in various plant organs and species. First, converging evidence is gathered from the literature that water deficit generally increases C concentration in plant organs. The hypothesis is raised that this could be due to organ expansion (as a major C sink) being affected earlier and more intensively than photosynthesis (C source) and metabolism. How such an increase is likely to interact with C signalling is not known. Hence, the literature is reviewed for possible links between C and stress signalling that could take part in this interaction. Finally, the possible impact of water deficit-induced C accumulation on growth is questioned for various sink organs of several species by combining published as well as new experimental data or data generated using a modelling approach. To this aim, robust correlations between C availability and sink organ growth are reported in the absence of water deficit. Under water deficit, relationships weaken or are modified suggesting release of the influence of C availability on sink organ growth. These results are interpreted as the signature of a transition from source to sink growth limitation under water deficit.     

Plant water deficits and insect responses: the preference of Corythucha arcuata (Heteroptera: Tingidae) for the foliage of white oak, Quercus alba

ABSTRACT.

• 1 The preference of Corythucha arcuata (Say) (Heteroptera: Tingidae) for the foliage of Quercus alba L. (Fagaceae) grown under water deficit was examined. Potted Q.alba saplings were grown under controlled levels of water deficit in a glasshouse. Adult C.arcuata were exposed to excised Q.alba foliage under controlled conditions in a growth chamber, and feeding preference assessed by recording the percentage of individuals feeding on leaves from each level of water deficit.
• 2 Two experiments were performed. In the first, all foliage was supplemented with water (via the petiole) to alleviate physical changes in the leaves that accompany plant water deficits. In the second experiment, leaves were not supplemented with water.
• 3 C.arcuata had no preference when presented water supplemented leaves, but preferred leaves that were fully hydrated when presented leaves that were not supplemented with water.
• 4 These results suggest that plant water deficits are important in determining the feeding preference of C.arcuata because they lead to avoidance of drought grown foliage. This avoidance may be caused by an increase in the metabolic cost to the herbivore of maintaining water balance when ingesting drought grown foliage or to an increase in foliage toughness.

     

Soil water deficits decrease the internal conductance to CO2 transfer but atmospheric water deficits do not

The internal conductance to CO₂ supply from substomatal cavitiesto sites of carboxylation poses a large limitation to photosynthesis.It is known that internal conductance is decreased by soil waterdeficits, but it is not known if it is affected by atmosphericwater deficits (i.e. leaf to air vapour pressure deficit, VPD).The aim of this paper was to examine the responses of internalconductance to atmospheric and soil water deficits in seedlingsof the evergreen perennial Eucalyptus regnans F. Muell and theherbaceous plants Solanum lycopersicum (formerly Lycopersiconesculentum) Mill. and Phaseolus vulgaris L. Internal conductancewas estimated with the variable J method from concurrent measurementsof gas exchange and fluorescence. In all three species steady-statestomatal conductance decreased by 30% as VPD increased from1 kPa to 2 kPa. In no species was internal conductance affectedby VPD despite large effects on stomatal conductance. In contrast,soil water deficits decreased stomatal conductance and internalconductance of all three species. Decreases in stomatal andinternal conductance under water deficit were proportional,but this proportionality differed among species, and thus therelationship between stomatal and internal conductance differedamong species. These findings indicate that soil water deficitsaffect internal conductance while atmospheric water deficitsdo not. The reasons for this distinction are unknown but areconsistent with soil and atmospheric water deficits having differingeffects on leaf physiology and/or root–shoot communication. Key words: Carbon dioxide, drought, internal conductance, mesophyll conductance, photosynthesis, stomatal conductance, transfer conductance, vapour pressure deficit, water deficit Received 11 October 2007; Revised 9 November 2007 Accepted 15 November 2007