Nitrogen enrichment lowers Betula pendula green and yellow leaf stoichiometry irrespective of effects of elevated carbon dioxide

Deep rooting has been identified as strategy for desiccation avoidance in natural vegetation as well as in crops like rice and sorghum. The objectives of this study were to determine root morphology and water uptake of four inbred lines of tropical maize (Zea mays L.) differing in their adaptation to drought. The specific questions were i) if drought tolerance was related to the vertical distribution of the roots, ii) whether root distribution was adaptive or constitutive, and iii) whether it affected water extraction, water status, and water use efficiency (WUE) of the plant. In the main experiment, seedlings were grown to the V5 stage in growth columns (0.80 m high) under well-watered (WW) and water-stressed (WS) conditions. The depth above which 95 % of all roots were located (D₉₅) was used to estimate rooting depth. It was generally greater for CML444 and Ac7729/TZSRW (P2) compared to SC-Malawi and Ac7643 (P1). The latter had more lateral roots, mainly in the upper part of the soil column. The increase in D₉₅ was accompanied by increases in transpiration, shoot dry weight, stomatal conductance and relative water content without adverse effects on the WUE. Differences in the morphology were confirmed in the V8 stage in large boxes: CML444 with thicker (0.14 mm) and longer (0.32 m) crown roots compared to SC-Malawi. Deep rooting, drought sensitive P2 showed markedly reduced WUE, likely due to an inefficient photosynthesis. The data suggest that a combination of high WUE and sufficient water acquisition by a deep root system can increase drought tolerance.

     

Increased contribution of wheat nocturnal transpiration to daily water use under drought

Increasing evidence suggests that in crops, nocturnal water use could represent 30% of daytime water consumption, particularly in semi‐arid and arid areas. This raises the questions of whether nocturnal transpiration rates (TR_N) are (1) less influenced by drought than daytime TR (TR_D), (2) increased by higher nocturnal vapor pressure deficit (VPD_N), which prevails in such environments and (3) involved in crop drought tolerance. In this investigation, we addressed those questions by subjecting two wheat genotypes differing in drought tolerance to progressive soil drying under two long‐term VPD_N regimes imposed under naturally fluctuating conditions. A first goal was to characterize the response curves of whole‐plant TR_N and TR_N/TR_D ratios to progressive soil drying. A second goal was to examine the effect of VPD_N increase on TR_N response to soil drying and on 13 other developmental traits. The study revealed that under drought, TR_N was not responsive to progressive soil drying and – intriguingly – that TR_N seemingly increased with drought under high VPD_N consistently for the drought‐sensitive genotype. Because TR_D was concomitantly decreasing with progressive drought, this resulted in TR_N representing up to 70% of TR_D at the end of the drydown. In addition, under drought, VPD_N increase was found not to influence traits such as leaf area or stomata density. Overall, those findings indicate that TR_N contribution to daily water use under drought might be much higher than previously thought, that it is controlled by specific mechanisms and that decreasing TR_N under drought might be a valuable trait for improving drought tolerance.     

Comparative Water Relations of Phaseolus vulgaris L. and Phaseolus acutifolius Gray

Leaf area expansion, dry weight, and water relations of Phaseolus vulgaris L. and P. acutifolius Gray were compared during a drying cycle in the greenhouse to understand the characteristics which contribute to the superior drought tolerance of P. acutifolius. Stomates of P. acutifolius closed at a much higher water potential than those of P. vulgaris, delaying dehydration of leaf tissue. P. acutifolius had a more deeply penetrating root system, which also contributes to its drought tolerance. Root-shoot ratios did not differ between the two species either under well watered or water stressed conditions. Leaf osmotic potential was also similar in the two species, with no apparent osmotic adjustment during water stress. These results indicate that P. acutifolius postpones dehydration and suggest that sensitive stomates and a deeply penetrating root system are characteristics which, if incorporated into cultivated beans, might increase their drought tolerance.     

Water uptake efficiency and above- and belowground biomass development of sweet sorghum and maize under different water regimes

Background and aims

Lately sweet sorghum (S) has attracted great interest as an alternative feedstock for biofuel production due to its high yielding potential and better adaptation to drought than maize (M). However, little is known about the response of newly developed sweet sorghum genotypes to water deficits, especially at the root level and its water uptake patterns. The objective of this study was to compare the water uptake capacity, growth and developmental characteristics at the root and canopy levels of a sweet sorghum hybrid (Sorghum bicolor cv. Sucro 506) with those of maize (Zea mays cv. PR32F73) at two water regimes.

Methods

The trial was setup in a total of 20 rhizotrons (1?m³), where calibrated soil moisture probes were installed for monitoring and adjusting the soil moisture content to 25% (well-watered, W) and 12% (drought stress, D).

Results

DS was able to sustain its physiological activity close to that of WS plants, while maize was not. The biomass production potential of DS was reduced about 38%, while in maize the reduction was 47%. The water use efficiency (WUE), however, was increased by 20% in sweet sorghum and reduced in 5% in maize. Moreover, in contrast to maize the root length density and water uptake capacity of DS was enhanced. Root water uptake efficiency in DM was sustained close to its potential, but not in sweet sorghum.

Conclusions

In summary, the better adaptation to drought of sweet sorghum is explained by increased WUE, sustained physiological activity and enlarged root system. It is also associated with a reduced water uptake efficiency compared to its control but maintained compared to maize.     

Root cortical burden influences drought tolerance in maize

Background and Aims

Root cortical aerenchyma (RCA) increases water and nutrient acquisition by reducing the metabolic costs of soil exploration. In this study the hypothesis was tested that living cortical area (LCA; transversal root cortical area minus aerenchyma area and intercellular air space) is a better predictor of root respiration, soil exploration and, therefore, drought tolerance than RCA formation or root diameter.

Methods

RCA, LCA, root respiration, root length and biomass loss in response to drought were evaluated in maize (Zea mays) recombinant inbred lines grown with adequate and suboptimal irrigation in soil mesocosms.

Key Results

Root respiration was highly correlated with LCA. LCA was a better predictor of root respiration than either RCA or root diameter. RCA reduced respiration of large-diameter roots. Since RCA and LCA varied in different parts of the root system, the effects of RCA and LCA on root length were complex. Greater crown-root LCA was associated with reduced crown-root length relative to total root length. Reduced LCA was associated with improved drought tolerance.

Conclusions

The results are consistent with the hypothesis that LCA is a driver of root metabolic costs and may therefore have adaptive significance for water acquisition in drying soil.     

The relations between drought susceptibility index based on grain yield (DSIGY) and key physiological seedling traits in maize and triticale genotypes

The physiological reasons for the differences in sensitivity of C₃ and C₄ plant species to environmental stresses have not been thoroughly explained. In this study the effects of drought stress on the growth and selected physiological traits were examined in the seedlings of 13 single cross maize (C₄ plant) hybrids and 11 spring triticale (C₃ plant) breeding lines and varieties differing in drought sensitivity. For plants in the seedling stage the results demonstrated a genetic variation in dry matter accumulation of shoots and roots (DW_S, DW_R), number (N) and length (L) of particular components (seminal, seminal adventitious, nodal) of the root system, membrane injury by soil drought (LI_D), osmotic and high temperature stress (LI_OS, LI_HT), water potential (ψ), water loss (WL), grain germination in osmotic stress (FG, PI), and seedling survival (SS). Seedlings grown under moderate soil drought showed a decrease in dry matter of the top parts and roots and a decrease in the length of seminal, seminal adventitious and nodal roots in comparison to seedlings grown in control conditions. The observed harmful effects of drought stress were more distinct in drought sensitive genotypes. Used in this paper drought susceptibility indexes (DSI_GY) were calculated in other experiment by determining the changes in grain yield (GY) under two soil moisture levels (irrigated and drought). The variation of DSI_GY for maize ranges from 0.381 to 0.650 and for triticale from 0.354 to 0.578. The correlations between DSI_GY and laboratory tests (LI, FG, SS) confirmed that they are good indicators of drought tolerance in plants. The highest values of genetic variation were observed in LI, DW_S, SS and WL and the lowest in the measurements of ψ FG, PI, L_S, L_SA and L_N. The correlation coefficients between LI_OS and LI_HT tests were, in most of the considered cases, statistically significant, which indicates that in maize and triticale the mechanisms of membrane injury caused by simulated drought or high temperature are physiologically similar. It can be concluded that an approach to the breeding of maize and triticale for drought tolerance using these tests can be implemented on the basis of separate selection for each trait or for all of them simultaneously. In that case, it would be necessary to determine the importance of the trait in relation to growth phase, drought timing and level, as well as its associations with morphological traits contributing to drought tolerance. The obtained values of the correlation coefficient between laboratory tests suggest that the same physiological traits may be applied as selection criteria in drought tolerance of maize and triticale genotypes.     

Soil textures rather than root hairs dominate water uptake and soil–plant hydraulics under drought

Although the role of root hairs (RHs) in nutrient uptake is well documented, their role in water uptake and drought tolerance remains controversial. Maize (Zea mays) wild-type and its hair-defective mutant (Mut; roothairless 3) were grown in two contrasting soil textures (sand and loam). We used a root pressure chamber to measure the relation between transpiration rate (E) and leaf xylem water potential (ψ_{leaf_x}) during soil drying. Our hypotheses were: (1) RHs extend root–soil contact and reduce the ψ_{leaf_x} decline at high E in dry soils; (2) the impact of RHs is more pronounced in sand; and (3) Muts partly compensate for lacking RHs by producing longer and/or thicker roots. The ψ_{leaf_x}(E) relation was linear in wet conditions and became nonlinear as the soils dried. This nonlinearity occurred more abruptly and at less negative matric potentials in sand (ca. −10 kPa) than in loam (ca. −100 kPa). At more negative soil matric potentials, soil hydraulic conductance became smaller than root hydraulic conductance in both soils. Both genotypes exhibited 1.7 times longer roots in loam, but 1.6 times thicker roots in sand. No differences were observed in the ψ_{leaf_x}(E) relation and active root length between the two genotypes. In maize, RHs had a minor contribution to soil–plant hydraulics in both soils and their putative role in water uptake was smaller than that reported for barley (Hordeum vulgare). These results suggest that the role of RHs cannot be easily generalized across species and soil textures affect the response of root hydraulics to soil drying.

Root hairs of maize do not show evident contribution to root growth, water uptake, and soil–plant hydraulics, whereas soil textures affect the response of root hydraulics to soil drying.     

Root metaxylem and architecture phenotypes integrate to regulate water use under drought stress

At the genus and species level, variation in root anatomy and architecture may interact to affect strategies of drought avoidance. To investigate this idea, root anatomy and architecture of the drought‐sensitive common bean (Phaseolus vulgaris) and drought‐adapted tepary bean (Phaseolus acutifolius) were analyzed in relation to water use under terminal drought. Intraspecific variation for metaxylem anatomy and axial conductance was found in the roots of both species. Genotypes with high‐conductance root metaxylem phenotypes acquired and transpired more water per unit leaf area, shoot mass, and root mass than genotypes with low‐conductance metaxylem phenotypes. Interspecific variation in root architecture and root depth was observed where P. acutifolius has a deeper distribution of root length than P. vulgaris. In the deeper‐rooted P. acutifolius, genotypes with high root conductance were better able to exploit deep soil water than genotypes with low root axial conductance. Contrastingly, in the shallower‐rooted P. vulgaris, genotypes with low root axial conductance had improved water status through conservation of soil moisture for sustained water capture later in the season. These results indicate that metaxylem morphology interacts with root system depth to determine a strategy of drought avoidance and illustrate synergism among architectural and anatomical phenotypes for root function.     

Effects of silicon on Zea mays plants exposed to water and oxygen deficiency

Effects of shoot and root supplementation with silicon on the response of Zea mays L. plants to matric water potential (Ψ_m) and oxygen deficiency (waterlogging) stresses were studied. The soil water limitation (Ψ_m) and oxygen deprivation significantly reduced shoot dry weight, chlorophyll (Chl) content, ascorbic acid content, as well as leaf relative water content. Both soil drying and waterlogging caused a significant increase in the leaf membrane injury by heat (51°C) and dehydration (40% PEG) stresses. The levels of lipid peroxidation (POL) and hydrogen peroxide (H₂O₂) content were increased by excess soil drying and oxygen deficiency. Supplementary silicon at 1.0 mM significantly increased Chl content and improved water status. Concentrations of H₂O₂, MDA, and proline and leaf membrane injury were significantly reduced by Si application. The reverse helds true for ascorbic acid. The results of this study indicate that application of silicon might improve growth attributes, effectively mitigate the adverse effect of drought and waterlogging, and increase tolerance of maize plants. The silicon-induced improvement of drought and anoxia tolerance was associated with the increase in oxidative defense abilities.     

Interspecific differences in root architecture among maize and triticale genotypes grown under drought,waterlogging and soil compaction

Environmental stresses (soil compaction, drought, waterlogging) cause changes in plants’ root system structure, also affecting the growth of above-ground parts. The aim of this study was to estimate phenotypic variation among maize and triticale genotypes in root penetration ability through petrolatum-wax-layer (RPA). Also, the effect of shortage or excess of soil water on dry matter of shoots and roots and morphological changes in root system structure in sensitive and resistant maize and triticale genotypes grown in low or high soil compaction level was evaluated. To estimate RPA index, the petrolatum-wax-layer method (PWL) was used. The strength of three petrolatum-wax concentrations 60, 50 and 40 % was 0.52, 1.07 and 1.58 MPa, respectively. High coefficients of variation (CV) were observed in 0.52 and 1.07 MPa and for maize were 19.2 and 21.7 %, and for triticale, 12.5 and 18.3 %, respectively. The data indicate that the use of PWL technique is an effective screening method, and makes it possible to divide the genotypes into resistant and sensitive groups. The second part of this study investigated a multistress effect of soil compaction combined with drought or waterlogging on root and shoot growth and morphological changes in root system structure of maize and triticale genotypes differing in susceptibility to environmental stresses. Seedlings were grown for 4 weeks in root-boxes under conditions of low (LSC 1.1 g cm^?3) or severe (SSC 1.6 g cm^?3) soil compaction. Drought or waterlogging stresses were applied for 2 weeks from 14th to 28th day. In comparison to LSC treatment, in SSC treatment the decrease in dry matter of shoots and roots was greater for sensitive genotypes of maize and triticale (Ancora, CHD-147). Soil drought or waterlogging caused greater decrease of dry matter of shoots and roots in seedlings grown in SSC in comparison to LSC. The root penetration index (RPI) was estimated as a ratio of root dry matter in 15–40 cm root-box layer to total root dry matter. On the basis of RPI it was possible to group the genotypes according to their ability to distribute roots in soil profile. In comparison to LSC, SSC exerted a strong influence on the length of seminal and seminal adventitious roots, as well as the number and length of L- and S-type lateral roots developed on seminal and nodal roots. In both species the restriction effect of soil compaction on number and length of roots was more severe in sensitive (Ankora, CHD-147) than in resistant (Tina, CHD-247) genotypes. The restriction in roots propagation was greater in triticale than in maize. Exposure to drought or waterlogging in the case of genotypes grown in LSC and SSC treatments caused a decrease in number and length of particular components of root system structure. In both species the decrease of root number and length in plants grown under waterlogging was greater than under drought. The observed changes in root system were greater in sensitive (Ankora, CHD147) than in resistant (Tina, CHD-247) genotypes. Statistically significant correlations were found between RPA and RPI and also between these indexes and soil compaction, drought and waterlogging susceptibility indexes. This indicates that genotypes resistant to soil compaction were resistant to drought or waterlogging and also that genotypes resistant to drought were resistant to waterlogging.     

Steep,cheap and deep: an ideotype to optimize water and N acquisition by maize root systems

Background

A hypothetical ideotype is presented to optimize water and N acquisition by maize root systems. The overall premise is that soil resource acquisition is optimized by the coincidence of root foraging and resource availability in time and space. Since water and nitrate enter deeper soil strata over time and are initially depleted in surface soil strata, root systems with rapid exploitation of deep soil would optimize water and N capture in most maize production environments.• The ideotype Specific phenes that may contribute to rooting depth in maize include (a) a large diameter primary root with few but long laterals and tolerance of cold soil temperatures, (b) many seminal roots with shallow growth angles, small diameter, many laterals, and long root hairs, or as an alternative, an intermediate number of seminal roots with steep growth angles, large diameter, and few laterals coupled with abundant lateral branching of the initial crown roots, (c) an intermediate number of crown roots with steep growth angles, and few but long laterals, (d) one whorl of brace roots of high occupancy, having a growth angle that is slightly shallower than the growth angle for crown roots, with few but long laterals, (e) low cortical respiratory burden created by abundant cortical aerenchyma, large cortical cell size, an optimal number of cells per cortical file, and accelerated cortical senescence, (f) unresponsiveness of lateral branching to localized resource availability, and (g) low K_m and high V_max for nitrate uptake. Some elements of this ideotype have experimental support, others are hypothetical. Despite differences in N distribution between low-input and commercial maize production, this ideotype is applicable to low-input systems because of the importance of deep rooting for water acquisition. Many features of this ideotype are relevant to other cereal root systems and more generally to root systems of dicotyledonous crops.     

Drought Responses of Foliar Metabolites in Three Maize Hybrids Differing in Water Stress Tolerance

Maize (Zea mays L.) hybrids varying in drought tolerance were treated with water stress in controlled environments. Experiments were performed during vegetative growth and water was withheld for 19 days beginning 17 days after sowing. Genotypic comparisons used measured changes of leaf water potential or results were expressed by time of treatment. Total dry matter of the drought tolerant hybrid on the final harvest was 53% less than that of the intermediate and susceptible maize hybrids when plants were water sufficient. This showed that maize hybrids selected for extreme drought tolerance possessed a dwarf phenotype that affected soil water contents and leaf water potentials. Changes of shoot and root growth, leaf water potential, net photosynthesis and stomatal conductance in response to the time of water stress treatment were diminished when comparing the drought tolerant to the intermediate or susceptible maize hybrids. Genotypic differences were observed in 26 of 40 total foliar metabolites during water stress treatments. Hierarchical clustering revealed that the tolerant maize hybrid initiated the accumulation of stress related metabolites at higher leaf water potentials than either the susceptible or intermediate hybrids. Opposite results occurred when changes of metabolites in maize leaves were expressed temporally. The above results demonstrated that genotypic differences were readily observed by comparing maize hybrids differing in drought tolerance based on either time of treatment or measured leaf water potential. Current findings provided new and potentially important insights into the mechanisms of drought tolerance in maize.     

The relationship between seedling growth and grain yield under drought conditions in maize and triticale genotypes

The effects of drought stress on seedlings?? growth and grain yield of 13 single cross maize hybrids and 11 breeding lines and cultivars of spring triticale were studied in greenhouse and field experiments. In the field experiment, the drought susceptibility index (DSI_GY) was calculated by determining the change in grain yield (GY) in conditions with two soil moisture levels (IR, irrigated; D, drought). In the greenhouse experiment the response to soil drought was evaluated using DSI_DW, by determining changes in the dry weight (DW) of vegetative plant parts. Marked variations in GY and DW were observed among the studied genotypes. In control conditions, the GY and DW in drought-sensitive genotypes were higher compared to the drought-resistant ones; but in drought conditions, the decreases in GY and DW in resistant genotypes were smaller than in drought-sensitive ones. DSI_GY and DSI_DW revealed variations in the degree of drought tolerance among the examined maize and triticale genotypes. The values of DSI_GY in the field experiment and DSI_DW in the greenhouse experiment enabled a division of the studied genotypes into drought-resistant or -sensitive groups. A close correlation between DSI_GY and DSI_DW was found. The positive linear correlation and determination coefficients between DSI_GY and DSI_DW were statistically significant (P?=?0.05), being equal to R ²?=?0.614 (maize) and R ²?=?0.535 (triticale). The ranking of the studied genotypes based on DSI_GY was in most cases consistent with the ranking based on DSI_DW, which indicates that genetically conditioned drought tolerance is similar for plants in the seedling and reproductive growth stages or may at least partly have a common genetic background.     

Aerenchyma development in different root zones of maize genotypes under water limitation and different phosphorus nutrition

Root cortical aerenchyma (RCA) is suggested to reduce metabolic cost for root growth, but it might lower water uptake by plants. The objective of this work was to evaluate the effects of drought and phosphorus on the RCA development along the root axis and to elucidate its role in water stress tolerance of two maize genotypes. Plants of drought-tolerant DKB390 and drought-sensitive BRS1010 genotypes were grown in Vermiculite at field capacity of 100, 75, 50, and 25 % and supplied with 0.1, 0.4, and 0.8 mM phosphorus. Growth parameters, RCA, and plant P content were evaluated for all plants. Higher RCA development was observed in DKB390 than in BRS1010. Drought reduced the percentage of RCA in the root-hair zone of both genotypes but increased its development in the root maturation zone. Phosphorus limitation enhanced RCA development only in the DKB390. Under drought stress, DKB390 showed resilient growth whereas growth was inhibited in BRS1010. Higher root P content was related to its higher supply. Therefore, RCA formation was induced either by drought or by phosphorus limitation, while no interaction was evident. The RCA development varied along the root axis in order to balance water and phosphorus uptake and the drought response was genotype dependent.     

The effect of drought stress on chlorophyll fluorescence in Lolium-Festuca hybrids

The effects of drought on photochemical efficiency of PSII in leaves of 22 hybrids of Festuca pratensis × Lolium multiflorum and Festuca pratensis × Lolium perenne and of Festuca pratensis cv. Skra were investigated. A significant decrease of electron transport efficiency (about 25%) in PSII (Φ_PSII) was not found before 9 days of seedling growth in hydroponics with water potential (Ψ_w) equal to −0.8 MPa (simulated “soil drought”). The decrease of Φ_PSII was similarly related to that of excitation energy capture by open PSII reaction centre (Fv’/Fm’) and also to the decrease of the proportion of oxidized to reduced Q_A (photochemical fluorescence quenching, q_p). According to the drought prolongation, variation of all parameters of fluorescence between genotypes significantly increased. The seedlings of some genotypes were able to recover electron transport efficiency in PSII after increasing water potential in nutrient solution (removing the “soil drought”). When plants grew in containers with soil and 4 genotypes with the highest sensitivity of electron transport to drought (S) as well as 4 genotypes with the highest tolerance (T) were compared 17 days after watering ceased, Ψ_w in leaves considerably decreased, but the differences between S and T genotypes were often not significant in this respect. The differences between S and T genotypes, as values of Fv/Fm were concerned, also appeared small (about 5%), similarly as that of Fv’/Fm’ (5%), q_p (12%) and Φ_PSII (about 15%). Drought stress increased non-photochemical quenching of chlorophyll fluorescence (NPQ) 15 to 47% and this could protect the PSII reaction centres from damages because of energy excess. The increase of NPQ was not closely connected with drought resistance of plants because it was similar in some genotypes tolerant to dehydration as well as in sensitive ones. The results of the experiments suggest that resources of genetic variability in Festulolium may be sufficient for revealing differences between genotypes on the basis of measurement of chlorophyll a fluorescence, as far as their tolerance to soil drought is concerned. As the tolerance of PSII against drought is high, the determinations of fluorescence should be performed rather under severe stress. Such methods seem to be useful for selection of genotypes with high drought tolerance as well as with the ability to at least partial repairing of PSII after drought.     

Root cortical aerenchyma improves the drought tolerance of maize (Zea mays L.)

Root cortical aerenchyma (RCA) reduces root respiration in maize by converting living cortical tissue to air volume. We hypothesized that RCA increases drought tolerance by reducing root metabolic costs, permitting greater root growth and water acquisition from drying soil. To test this hypothesis, recombinant inbred lines with high and low RCA were observed under water stress in the field and in soil mesocosms in a greenhouse. In the field, lines with high RCA had 30% more shoot biomass at flowering compared with lines with low RCA under water stress. Root length density in deep soil was significantly greater in the high RCA lines compared with the low RCA lines. Mid‐day leaf relative water content in the high RCA lines was 10% greater than in the low RCA lines under water stress. The high RCA lines averaged eight times the yield of the low RCA lines under water stress. In mesocosms, high RCA lines had less seminal root respiration, deeper rooting, and greater shoot biomass compared with low RCA lines under water stress. These results support the hypothesis that RCA is beneficial for drought tolerance in maize by reducing the metabolic cost of soil exploration.     

Root hairs are the most important root trait for rhizosheath formation of barley (Hordeum vulgare), maize (Zea mays) and Lotus japonicus (Gifu)

Background and AimsRhizosheaths are defined as the soil adhering to the root system after it is extracted from the ground. Root hairs and mucilage (root exudates) are key root traits involved in rhizosheath formation, but to better understand the mechanisms involved their relative contributions should be distinguished.MethodsThe ability of three species [barley (Hordeum vulgare), maize (Zea mays) and Lotus japonicus (Gifu)] to form a rhizosheath in a sandy loam soil was compared with that of their root-hairless mutants [bald root barley (brb), maize root hairless 3 (rth3) and root hairless 1 (Ljrhl1)]. Root hair traits (length and density) of wild-type (WT) barley and maize were compared along with exudate adhesiveness of both barley and maize genotypes. Furthermore, root hair traits and exudate adhesiveness from different root types (axile versus lateral) were compared within the cereal species.Key ResultsPer unit root length, rhizosheath size diminished in the order of barley > L. japonicus > maize in WT plants. Root hairs significantly increased rhizosheath formation of all species (3.9-, 3.2- and 1.8-fold for barley, L. japonicus and maize, respectively) but there was no consistent genotypic effect on exudate adhesiveness in the cereals. While brb exudates were more and rth3 exudates were less adhesive than their respective WTs, maize rth3 bound more soil than barley brb. Although both maize genotypes produced significantly more adhesive exudate than the barley genotypes, root hair development of WT barley was more extensive than that of WT maize. Thus, the greater density of longer root hairs in WT barley bound more soil than WT maize. Root type did not seem to affect rhizosheath formation, unless these types differed in root length.ConclusionsWhen root hairs were present, greater root hair development better facilitated rhizosheath formation than root exudate adhesiveness. However, when root hairs were absent root exudate adhesiveness was a more dominant trait.     

Reduced Root Cortical Cell File Number Improves Drought Tolerance in Maize

We tested the hypothesis that reduced root cortical cell file number (CCFN) would improve drought tolerance in maize (Zea mays) by reducing the metabolic costs of soil exploration. Maize genotypes with contrasting CCFN were grown under well-watered and water-stressed conditions in greenhouse mesocosms and in the field in the United States and Malawi. CCFN ranged from six to 19 among maize genotypes. In mesocosms, reduced CCFN was correlated with 57% reduction of root respiration per unit of root length. Under water stress in the mesocosms, genotypes with reduced CCFN had between 15% and 60% deeper rooting, 78% greater stomatal conductance, 36% greater leaf CO₂ assimilation, and between 52% to 139% greater shoot biomass than genotypes with many cell files. Under water stress in the field, genotypes with reduced CCFN had between 33% and 40% deeper rooting, 28% lighter stem water oxygen isotope enrichment (δ¹⁸O) signature signifying deeper water capture, between 10% and 35% greater leaf relative water content, between 35% and 70% greater shoot biomass at flowering, and between 33% and 114% greater yield than genotypes with many cell files. These results support the hypothesis that reduced CCFN improves drought tolerance by reducing the metabolic costs of soil exploration, enabling deeper soil exploration, greater water acquisition, and improved growth and yield under water stress. The large genetic variation for CCFN in maize germplasm suggests that CCFN merits attention as a breeding target to improve the drought tolerance of maize and possibly other cereal crops.Drought is a primary constraint to global crop production (Schmidhuber and Tubiello, 2007), and global climate change is likely to increase the risk of drought, especially in rain-fed agriculture (Battisti and Naylor, 2009; Burke et al., 2009; Mishra and Cherkauer, 2010; Lobell et al., 2011). Therefore, the development of crops with greater drought tolerance is an important global objective. Yield under drought is often not an efficient selection criterion in drought breeding programs, since yield is affected by many elements of the phenotype and the environment, interacting in complex and often unknown ways. Trait-based selection or ideotype breeding is generally a more efficient selection strategy, permitting the identification of useful sources of variation among lines that have poor agronomic adaptation, elucidation of genotype-by-environment interactions, and informed trait stacking (Araus et al., 2002, 2008; Manschadi et al., 2006; Lynch, 2007b, 2011; York et al., 2013).In most agroecosystems, the topsoil dries before the subsoil as drought progresses. In such environments, plants with deeper roots are able to acquire water available in deeper soil domains that may not be available to plants with shallower roots (Ludlow and Muchow, 1990; Ho et al., 2005; Hammer et al., 2009). An ideotype has been proposed to guide the breeding of crops with deeper roots and, therefore, greater water acquisition from drying soil, called Steep, Cheap, and Deep, integrating architectural, anatomical, and physiological phenes (Lynch, 2013). The term Cheap denotes phenes that reduce the metabolic cost of soil exploration, which is an important limitation to the acquisition of scarce soil resources, including water in dry soil (Fan et al., 2003; Lynch, 2007b; Zhu et al., 2010; Postma and Lynch, 2011a, 2011b; Jaramillo et al., 2013). Plant resource allocation to root growth typically increases under drought to enhance water acquisition; therefore, the metabolic cost of root growth becomes a significant component of plant fitness and adaptation under drought (Lynch, 2007b, 2013). Therefore, a plant that is able to access water in deep soil domains at reduced metabolic cost will have superior productivity, because it will have more metabolic resources available for further resource acquisition, growth, and reproduction. Evidence in support of this hypothesis comes from empirical and modeling studies for maize (Zea mays) under water and edaphic stress (Lynch, 2007a; Zhu et al., 2010; Postma and Lynch, 2011a, 2011b; Jaramillo et al., 2013).Root cortical aerenchyma (RCA) is the enlarged air space in the root cortex that forms either through cell death or cell separation (Evans, 2004). RCA is associated with a disproportionate reduction of root respiration in maize by converting living cortical tissue to air volume (Fan et al., 2003; Zhu et al., 2010). Reduction of root metabolic costs permits more internal resources to be allocated to greater root growth and, consequently, greater soil resource acquisition. RCA formation is also associated with a reduction of phosphorus content in root tissue on a volume basis, since air spaces do not contain phosphorus (Fan et al., 2003), and with improved growth in low-phosphorus soil (Lynch, 2011). RCA also reduces the nitrogen content of root tissue and is beneficial for nitrogen capture and maize growth on low-nitrogen soils (Saengwilai, 2014a). Modeling studies suggest that RCA improves crop adaptation to suboptimal nutrient availability by reducing the metabolic costs of soil exploration (Postma and Lynch, 2011a, 2011b). Under drought, Zhu et al. (2010) found that maize genotypes with more RCA had five times greater biomass and eight times greater yield than genotypes with less RCA. Living cortical area (LCA) is total transverse root cortical area minus RCA area. Jaramillo et al. (2013) found that root respiration is positively correlated with LCA, and a 3.5-fold reduction in LCA is associated with a 2.5-fold improvement in plant growth under drought. These results indicate that the metabolic demand of living cortical tissue is a primary determinant of root growth, soil exploration, and resource acquisition in soil environments with suboptimal resource availability.This study builds on earlier studies indicating that substantial reduction of root metabolic cost is associated with variation in LCA. The cortex of the maize root is composed of several concentric layers of parenchyma cells, the number of which we refer to as the cortical cell file number (CCFN). Recently, Burton et al. (2013) reported that there is 3-fold variation for CCFN in Zea spp. In that study, the variation was wider in maize landraces (six to 16 cell files) than in wild Zea spp. (seven to 13 cell files). It has been proposed that reduced CCFN would decrease the metabolic costs of root growth and maintenance, in terms of both the carbon cost of root respiration and the nutrient content of living tissue, by reducing the proportion of root volume occupied by living cortical tissue, which has greater metabolic demands than the stele (Lynch, 2013). However, the physiological utility of CCFN has not been explored.The objective of this study was to test the hypothesis that reduced CCFN would reduce root respiration, permitting greater rooting depth, thereby enhancing water acquisition and improving both plant growth and yield under water stress.     

Significance of root hairs for plant performance under contrasting field conditions and water deficit

Background and AimsPrevious laboratory studies have suggested selection for root hair traits in future crop breeding to improve resource use efficiency and stress tolerance. However, data on the interplay between root hairs and open-field systems, under contrasting soils and climate conditions, are limited. As such, this study aims to experimentally elucidate some of the impacts that root hairs have on plant performance on a field scale.MethodsA field experiment was set up in Scotland for two consecutive years, under contrasting climate conditions and different soil textures (i.e. clay loam vs. sandy loam). Five barley (Hordeum vulgare) genotypes exhibiting variation in root hair length and density were used in the study. Root hair length, density and rhizosheath weight were measured at several growth stages, as well as shoot biomass, plant water status, shoot phosphorus (P) accumulation and grain yield.Key ResultsMeasurements of root hair density, length and its correlation with rhizosheath weight highlighted trait robustness in the field under variable environmental conditions, although significant variations were found between soil textures as the growing season progressed. Root hairs did not confer a notable advantage to barley under optimal conditions, but under soil water deficit root hairs enhanced plant water status and stress tolerance resulting in a less negative leaf water potential and lower leaf abscisic acid concentration, while promoting shoot P accumulation. Furthermore, the presence of root hairs did not decrease yield under optimal conditions, while root hairs enhanced yield stability under drought.ConclusionsSelecting for beneficial root hair traits can enhance yield stability without diminishing yield potential, overcoming the breeder’s dilemma of trying to simultaneously enhance both productivity and resilience. Therefore, the maintenance or enhancement of root hairs can represent a key trait for breeding the next generation of crops for improved drought tolerance in relation to climate change.