Andreas J. Wild
Current Organizations / Institutes
The organizations/institutes from the scholar's latest article
Research Subjects
Andreas J. Wild has a total of 36 co-authors. Here are the top co-authors by number of shared publications:
- Johanna Pausch
Agroecology, Bayreuth Center of Ecology and Environmental Research (BayCEER), University of Bayreuth, Bayreuth, Germany
9 shared publications - Carsten Werner Mueller
Institute of Ecology Technische Universität Berlin Berlin Germany
6 shared publications - Franziska Steiner
Soil Science, TUM School of Life Sciences, Technical University of Munich , Freising
6 shared publications - Nicolas Tyborski
Ecological Microbiology, Bayreuth Center of Ecology and Environmental Research (BayCEER), University of Bayreuth , DE-95448 Bayreuth
6 shared publications - Shu-Yin Tung
Institute for Agroecology and Organic Farming, Bavarian State Research Centre for Agriculture (LfL) , DE-85354 Freising
6 shared publications - Alix Vidal
Soil Biology group, Wageningen University and Research, Wageningen, AA 6700, Netherlands
5 shared publications
Latest Publications
- Open Access
- 17 May 2026
- Published by Elsevier in Soil Biology and Biochemistry
- Vol. 220
Plant, mycorrhizal and saprotrophic fungal contributions to soil organic matter fractions differ between tree-mycorrhizal types
- 17 May 2026
- Published by Elsevier in Soil Biology and Biochemistry
- Vol. 220
Open AccessPlant, mycorrhizal and saprotrophic fungal contributions to soil organic matter fractions differ between tree-mycorrhizal types
- Open Access
- 14 January 2026
- Published by Oxford University Press (OUP) in ISME Communications
- Vol. 6 (1)
Breeding effects on the root-associated microbiome of Zea mays L. are linked to plant-induced variation in soil water potentials
Modern crop varieties may exert reduced influence on their microbiome compared to their progenitors, as plant-microbe interactions were not targeted during breeding. Moreover, formerly beneficial microbiome functions might no longer be relevant in modern agricultural ecosystems. We hypothesised that such patterns could become particularly evident under drought, since drought-tolerance has not been a primary breeding target. To test this, we grew six maize landraces (released before 1945) and six modern varieties (released from 2010 onwards) in a field under ambient and 60% reduced precipitation. The experiment was repeated over two years, differing in amounts and temporal distributions of precipitation. We assessed the composition of root-associated prokaryotic communities during grain filling by 16S rRNA gene metabarcoding. Intra-variety dispersion in microbiome composition relative to plant biomass-based dispersion was higher in modern varieties, suggesting breeding may have affected plant control over microbiomes. Besides that, shifts in microbiome composition between landraces and modern varieties were driven mainly by the plants' impact on soil water potentials. Consequently, the taxa that increased in relative abundance during soil drying, mainly Actinomycetota, were similar between landraces and modern varieties. Exploring microbiome-mediated alleviation of drought effects, therefore, appears promising also for applications in modern agricultural ecosystems. Specifically, filamentous Streptomyces spp. potentially contributed to soil aggregate stability, which should be further investigated in the context of drought mitigation. The reduced plant control over microbiome composition of modern varieties suggested by dispersion analysis likely has functional implications beyond microbiome adaptation to drought and should be considered in future assessments of breeding.
- 14 January 2026
- Published by Oxford University Press (OUP) in ISME Communications
- Vol. 6 (1)
Open AccessBreeding effects on the root-associated microbiome of Zea mays L. are linked to plant-induced variation in soil water potentials
Modern crop varieties may exert reduced influence on their microbiome compared to their progenitors, as plant-microbe interactions were not targeted during breeding. Moreover, formerly beneficial microbiome functions might no longer be relevant in modern agricultural ecosystems. We hypothesised that such patterns could become particularly evident under drought, since drought-tolerance has not been a primary breeding target. To test this, we grew six maize landraces (released before 1945) and six modern varieties (released from 2010 onwards) in a field under ambient and 60% reduced precipitation. The experiment was repeated over two years, differing in amounts and temporal distributions of precipitation. We assessed the composition of root-associated prokaryotic communities during grain filling by 16S rRNA gene metabarcoding. Intra-variety dispersion in microbiome composition relative to plant biomass-based dispersion was higher in modern varieties, suggesting breeding may have affected plant control over microbiomes. Besides that, shifts in microbiome composition between landraces and modern varieties were driven mainly by the plants' impact on soil water potentials. Consequently, the taxa that increased in relative abundance during soil drying, mainly Actinomycetota, were similar between landraces and modern varieties. Exploring microbiome-mediated alleviation of drought effects, therefore, appears promising also for applications in modern agricultural ecosystems. Specifically, filamentous Streptomyces spp. potentially contributed to soil aggregate stability, which should be further investigated in the context of drought mitigation. The reduced plant control over microbiome composition of modern varieties suggested by dispersion analysis likely has functional implications beyond microbiome adaptation to drought and should be considered in future assessments of breeding.
- Open Access
- 21 April 2025
- Published by Springer Nature in Plant and Soil
- Vol. 514 (1) , 1241-1261
Soil drying shapes rhizosheath properties and their link with maize yields across different soils
Background and Aims: Biophysicochemical soil properties in the rhizosheath are pivotal for crop yields and drive organic carbon cycling in agricultural soils. Yet, it remains uncertain how moderate soil drought may alter and interfere with rhizosheath properties in diverse soil types, and whether specific rhizosheath traits benefit crop yields under different water availability in heterogeneous field environments. Methods: Maize ( Zea mays L.) was grown under ambient precipitation and moderate drought (60% precipitation exclusion) at two field sites differing in physicochemical soil properties, such as texture and amounts of soil organic matter (SOM). Rhizosheath properties, namely soil aggregation as well as content and distribution of carbon (C) and nitrogen (N), were analyzed and, in conjunction with root traits, related to maize yields. Results: Under moderate soil drought, net rhizosheath-C concentrated spatially within the smaller rhizosheath in the form of more carbon-rich rhizodeposits. These effects were mediated by native soil properties, with rhizosheath structural stability decreasing stronger under drought in the finer-textured soil, allocating greater proportions of C and N to microaggregates. Rhizosheath and root properties were associated with maize yields. Yet, the influence and importance of belowground traits for crop yields varied with environmental conditions (soil x precipitation). Conclusion: The responsiveness of rhizosheath properties to moderate soil drought may influence the fate and turnover of root-derived C, and thus the overall drought resilience of SOM in agricultural soils. Moreover, our findings underscore the importance of rhizosheath properties for crop yields, yet highlighting that these relationships differ among environmental scenarios.
- 21 April 2025
- Published by Springer Nature in Plant and Soil
- Vol. 514 (1) , 1241-1261
Open AccessSoil drying shapes rhizosheath properties and their link with maize yields across different soils
Background and Aims: Biophysicochemical soil properties in the rhizosheath are pivotal for crop yields and drive organic carbon cycling in agricultural soils. Yet, it remains uncertain how moderate soil drought may alter and interfere with rhizosheath properties in diverse soil types, and whether specific rhizosheath traits benefit crop yields under different water availability in heterogeneous field environments. Methods: Maize ( Zea mays L.) was grown under ambient precipitation and moderate drought (60% precipitation exclusion) at two field sites differing in physicochemical soil properties, such as texture and amounts of soil organic matter (SOM). Rhizosheath properties, namely soil aggregation as well as content and distribution of carbon (C) and nitrogen (N), were analyzed and, in conjunction with root traits, related to maize yields. Results: Under moderate soil drought, net rhizosheath-C concentrated spatially within the smaller rhizosheath in the form of more carbon-rich rhizodeposits. These effects were mediated by native soil properties, with rhizosheath structural stability decreasing stronger under drought in the finer-textured soil, allocating greater proportions of C and N to microaggregates. Rhizosheath and root properties were associated with maize yields. Yet, the influence and importance of belowground traits for crop yields varied with environmental conditions (soil x precipitation). Conclusion: The responsiveness of rhizosheath properties to moderate soil drought may influence the fate and turnover of root-derived C, and thus the overall drought resilience of SOM in agricultural soils. Moreover, our findings underscore the importance of rhizosheath properties for crop yields, yet highlighting that these relationships differ among environmental scenarios.
- Open Access
- 1 November 2024
- Published by Wiley in GCB Bioenergy
- Vol. 16 (12)
Moderate Drought Constrains Crop Growth Without Altering Soil Organic Carbon Dynamics in Perennial Cup‐Plant and Silage Maize
Silage maize (Zea mays L.) intensification to maximise biomass production increases greenhouse gas emissions, accelerates climate change and intensifies the search for alternative bioenergy crops with high carbon (C) sequestration capacity. The perennial cup‐plant (Silphium perfoliatum L.) not only serves as a viable bioenergy source but may also be a promising soil C conservator. However, the dynamics of soil organic C (SOC) under the C3 cup‐plant, exposed to moderate drought conditions, that reduces growth rate without causing crop failure, compared with the drought‐tolerant C4 maize, remains unexplored. Here, we investigated in a lysimeter experiment the effects of moderate drought stress on crop growth and soil CO2 efflux under cup‐plant and silage maize compared with well‐watered conditions. Soil CO2 efflux along with root and shoot biomass, soil moisture and temperature as well as SOC and nitrogen (N) were measured over three consecutive years. Irrespective of the watering regime, cup‐plant induced a greater soil CO2 efflux (16% and 23% for 2020 and 2021, respectively), which was associated with higher root and shoot biomass compared with silage maize suggesting a substantial contribution of the roots to total soil CO2 efflux. In addition, soil CO2 efflux correlated negatively with soil dissolved N and positively with microbial C:N imbalance suggesting that low soil N availability influences soil CO2 efflux through processes related to N‐limitation such as N‐mining. Strikingly, moderate drought had no effect on soil CO2 efflux and C content and microbial biomass C, but increased dissolved organic C and microbial biomass N in both crops suggesting a complex interplay between C availability, N‐limitation and microbial adaptation under these conditions. Although cup‐plant increased soil CO2 efflux, the observed higher root and shoot biomass even under moderate drought conditions suggests a similar soil C management as silage maize; however, this still requires longer‐term investigation.
- 1 November 2024
- Published by Wiley in GCB Bioenergy
- Vol. 16 (12)
Open AccessModerate Drought Constrains Crop Growth Without Altering Soil Organic Carbon Dynamics in Perennial Cup‐Plant and Silage Maize
Silage maize (Zea mays L.) intensification to maximise biomass production increases greenhouse gas emissions, accelerates climate change and intensifies the search for alternative bioenergy crops with high carbon (C) sequestration capacity. The perennial cup‐plant (Silphium perfoliatum L.) not only serves as a viable bioenergy source but may also be a promising soil C conservator. However, the dynamics of soil organic C (SOC) under the C3 cup‐plant, exposed to moderate drought conditions, that reduces growth rate without causing crop failure, compared with the drought‐tolerant C4 maize, remains unexplored. Here, we investigated in a lysimeter experiment the effects of moderate drought stress on crop growth and soil CO2 efflux under cup‐plant and silage maize compared with well‐watered conditions. Soil CO2 efflux along with root and shoot biomass, soil moisture and temperature as well as SOC and nitrogen (N) were measured over three consecutive years. Irrespective of the watering regime, cup‐plant induced a greater soil CO2 efflux (16% and 23% for 2020 and 2021, respectively), which was associated with higher root and shoot biomass compared with silage maize suggesting a substantial contribution of the roots to total soil CO2 efflux. In addition, soil CO2 efflux correlated negatively with soil dissolved N and positively with microbial C:N imbalance suggesting that low soil N availability influences soil CO2 efflux through processes related to N‐limitation such as N‐mining. Strikingly, moderate drought had no effect on soil CO2 efflux and C content and microbial biomass C, but increased dissolved organic C and microbial biomass N in both crops suggesting a complex interplay between C availability, N‐limitation and microbial adaptation under these conditions. Although cup‐plant increased soil CO2 efflux, the observed higher root and shoot biomass even under moderate drought conditions suggests a similar soil C management as silage maize; however, this still requires longer‐term investigation.
- Open Access
- 22 March 2024
- Published by Wiley in Plant, Cell & Environment
- Vol. 47 (7) , 2524-2539
Unraveling root and rhizosphere traits in temperate maize landraces and modern cultivars: Implications for soil resource acquisition and drought adaptation
A holistic understanding of plant strategies to acquire soil resources is pivotal in achieving sustainable food security. However, we lack knowledge about variety‐specific root and rhizosphere traits for resource acquisition, their plasticity and adaptation to drought. We conducted a greenhouse experiment to phenotype root and rhizosphere traits (mean root diameter [Root D], specific root length [SRL], root tissue density, root nitrogen content, specific rhizosheath mass [SRM], arbuscular mycorrhizal fungi [AMF] colonization) of 16 landraces and 22 modern cultivars of temperate maize (Zea mays L.). Our results demonstrate that landraces and modern cultivars diverge in their root and rhizosphere traits. Although landraces follow a ‘do‐it‐yourself’ strategy with high SRLs, modern cultivars exhibit an ‘outsourcing’ strategy with increased mean Root Ds and a tendency towards increased root colonization by AMF. We further identified that SRM indicates an ‘outsourcing’ strategy. Additionally, landraces were more drought‐responsive compared to modern cultivars based on multitrait response indices. We suggest that breeding leads to distinct resource acquisition strategies between temperate maize varieties. Future breeding efforts should increasingly target root and rhizosphere economics, with SRM serving as a valuable proxy for identifying varieties employing an outsourcing resource acquisition strategy.
- 22 March 2024
- Published by Wiley in Plant, Cell & Environment
- Vol. 47 (7) , 2524-2539
Open AccessUnraveling root and rhizosphere traits in temperate maize landraces and modern cultivars: Implications for soil resource acquisition and drought adaptation
A holistic understanding of plant strategies to acquire soil resources is pivotal in achieving sustainable food security. However, we lack knowledge about variety‐specific root and rhizosphere traits for resource acquisition, their plasticity and adaptation to drought. We conducted a greenhouse experiment to phenotype root and rhizosphere traits (mean root diameter [Root D], specific root length [SRL], root tissue density, root nitrogen content, specific rhizosheath mass [SRM], arbuscular mycorrhizal fungi [AMF] colonization) of 16 landraces and 22 modern cultivars of temperate maize (Zea mays L.). Our results demonstrate that landraces and modern cultivars diverge in their root and rhizosphere traits. Although landraces follow a ‘do‐it‐yourself’ strategy with high SRLs, modern cultivars exhibit an ‘outsourcing’ strategy with increased mean Root Ds and a tendency towards increased root colonization by AMF. We further identified that SRM indicates an ‘outsourcing’ strategy. Additionally, landraces were more drought‐responsive compared to modern cultivars based on multitrait response indices. We suggest that breeding leads to distinct resource acquisition strategies between temperate maize varieties. Future breeding efforts should increasingly target root and rhizosphere economics, with SRM serving as a valuable proxy for identifying varieties employing an outsourcing resource acquisition strategy.
- Open Access
- 28 February 2024
- Published by Wiley in New Phytologist
- Vol. 242 (2) , 479-492
Rhizosheath drought responsiveness is variety‐specific and a key component of belowground plant adaptation
Summary: Biophysicochemical rhizosheath properties play a vital role in plant drought adaptation. However, their integration into the framework of plant drought response is hampered by incomplete mechanistic understanding of their drought responsiveness and unknown linkage to intraspecific plant–soil drought reactions. Thirty‐eight Zea mays varieties were grown under well‐watered and drought conditions to assess the drought responsiveness of rhizosheath properties, such as soil aggregation, rhizosheath mass, net‐rhizodeposition, and soil organic carbon distribution. Additionally, explanatory traits, including functional plant trait adaptations and changes in soil enzyme activities, were measured. Drought restricted soil structure formation in the rhizosheath and shifted plant–carbon from litter‐derived organic matter in macroaggregates to microbially processed compounds in microaggregates. Variety‐specific functional trait modifications determined variations in rhizosheath drought responsiveness. Drought responses of the plant–soil system ranged among varieties from maintaining plant–microbial interactions in the rhizosheath through accumulation of rhizodeposits, to preserving rhizosheath soil structure while increasing soil exploration through enhanced root elongation. Drought‐induced alterations at the root–soil interface may hold crucial implications for ecosystem resilience in a changing climate. Our findings highlight that rhizosheath soil properties are an intrinsic component of plant drought response, emphasizing the need for a holistic concept of plant–soil systems in future research on plant drought adaptation.
- 28 February 2024
- Published by Wiley in New Phytologist
- Vol. 242 (2) , 479-492
Open AccessRhizosheath drought responsiveness is variety‐specific and a key component of belowground plant adaptation
Summary: Biophysicochemical rhizosheath properties play a vital role in plant drought adaptation. However, their integration into the framework of plant drought response is hampered by incomplete mechanistic understanding of their drought responsiveness and unknown linkage to intraspecific plant–soil drought reactions. Thirty‐eight Zea mays varieties were grown under well‐watered and drought conditions to assess the drought responsiveness of rhizosheath properties, such as soil aggregation, rhizosheath mass, net‐rhizodeposition, and soil organic carbon distribution. Additionally, explanatory traits, including functional plant trait adaptations and changes in soil enzyme activities, were measured. Drought restricted soil structure formation in the rhizosheath and shifted plant–carbon from litter‐derived organic matter in macroaggregates to microbially processed compounds in microaggregates. Variety‐specific functional trait modifications determined variations in rhizosheath drought responsiveness. Drought responses of the plant–soil system ranged among varieties from maintaining plant–microbial interactions in the rhizosheath through accumulation of rhizodeposits, to preserving rhizosheath soil structure while increasing soil exploration through enhanced root elongation. Drought‐induced alterations at the root–soil interface may hold crucial implications for ecosystem resilience in a changing climate. Our findings highlight that rhizosheath soil properties are an intrinsic component of plant drought response, emphasizing the need for a holistic concept of plant–soil systems in future research on plant drought adaptation.
- Open Access
- 27 March 2022
- Published by Copernicus GmbH
High-throughput phenotyping of 38 maize varieties for the study of rhizosphere traits affecting agronomic resilience under drought stress
The occurrence of drought is likely to increase and intensify as a result of climate change, which poses a great challenge to agriculture. It is thus crucial to enhance agronomic resilience to secure food and feed production. Roots and root functioning as well as the interplay of roots with the surrounding soil, the rhizosphere, plays a key role in water acquisition of plants. Investigating rhizosphere traits is hence promising to shed light on future crops that better adapt to drought stress. A great strength of this study is the screening of various varieties which is facilitated by the high-throughput phenotyping method. It allows a wider coverage of traits and especially the genetic and phenetic diversities preserved in landraces.Maize (Zea mays L.), being one of the major cereal crops worldwide, was selected as the plant of study. A total of 38 varieties, which encompasses hybrid varieties, open pollinated varieties, and landraces, were screened in the “Moving Fields”, a greenhouse equipped with the high-throughput phenotyping facility in the Bavarian State Research Center for Agriculture. Maize plants were grown in mesocosms filled with loamy soil. Plants were exposed to two water treatments, well-watered and drought-stressed, during vegetative stem extension stage. Dynamic plant development was captured by continuous image acquisition. A visible light (RGB) camera was used to document the size and architecture of shoots and roots, while a chlorophyll fluorescence camera recorded the metabolic activity of shoots.Using shoot images, we compared variety-specific plant growth curves under well-watered and drought-stressed conditions to highlight the growth strategy of plants towards drought stress. The results reveal differences in growth inhibition during drought across varieties. In addition, differences in shoot and root dry weights are found between landraces and modern varieties. More analyses are in progress in search of rhizosphere traits and their influences on agronomic resilience.
- 27 March 2022
- Published by Copernicus GmbH
Open AccessHigh-throughput phenotyping of 38 maize varieties for the study of rhizosphere traits affecting agronomic resilience under drought stress
The occurrence of drought is likely to increase and intensify as a result of climate change, which poses a great challenge to agriculture. It is thus crucial to enhance agronomic resilience to secure food and feed production. Roots and root functioning as well as the interplay of roots with the surrounding soil, the rhizosphere, plays a key role in water acquisition of plants. Investigating rhizosphere traits is hence promising to shed light on future crops that better adapt to drought stress. A great strength of this study is the screening of various varieties which is facilitated by the high-throughput phenotyping method. It allows a wider coverage of traits and especially the genetic and phenetic diversities preserved in landraces.Maize (Zea mays L.), being one of the major cereal crops worldwide, was selected as the plant of study. A total of 38 varieties, which encompasses hybrid varieties, open pollinated varieties, and landraces, were screened in the “Moving Fields”, a greenhouse equipped with the high-throughput phenotyping facility in the Bavarian State Research Center for Agriculture. Maize plants were grown in mesocosms filled with loamy soil. Plants were exposed to two water treatments, well-watered and drought-stressed, during vegetative stem extension stage. Dynamic plant development was captured by continuous image acquisition. A visible light (RGB) camera was used to document the size and architecture of shoots and roots, while a chlorophyll fluorescence camera recorded the metabolic activity of shoots.Using shoot images, we compared variety-specific plant growth curves under well-watered and drought-stressed conditions to highlight the growth strategy of plants towards drought stress. The results reveal differences in growth inhibition during drought across varieties. In addition, differences in shoot and root dry weights are found between landraces and modern varieties. More analyses are in progress in search of rhizosphere traits and their influences on agronomic resilience.
- Open Access
- 27 March 2022
- Published by Copernicus GmbH
Responsiveness of maize to soil drying is related to a decrease in belowground hydraulic conductivity
Limited water supply is one of the largest impediments to food production worldwide in the light of climate change and increasing food demand. Stomatal regulation allows plants to promptly react to water stress and regulate water use. Although the coordination between stomatal closure and aboveground hydraulics has extensively been studied, our understanding of the impact of belowground hydraulics on stomatal regulation remains, as yet, incomplete. The overall objective of this study was to investigate the impact of belowground hydraulic conductivity as affected by differences in expressions of root and rhizosphere traits on the water use regulation of different maize genotypes.We have utilized a novel phenotyping facility to investigate the response of a selection of 48 maize (Zea mays L.) genotypes exhibiting different root and rhizosphere traits to soil drying. We measured the relation between leaf water potential, soil water potential, soil water content and transpiration rate, as well as root and rhizosphere traits (e.g. root length, rhizosheath mass) between genotypes. Our hypothesis is that stomatal response to soil drying is related to a loss in soil hydraulic conductivity and that key root and rhizosphere hydraulic traits affect such relation.We found that the genotypes differed in their responsiveness to drought and that such differences were related to belowground hydraulic traits. The critical soil water content at which plants started to decrease transpiration was related to a combination of plant- and rhizosphere traits (namely plant hydraulic conductance, maximum transpiration rate, root length and rhizosheath mass). Genotypes with a higher maximum transpiration rate and a higher plant hydraulic conductance and a smaller root system closed stomata in wetter soil conditions, meaning earlier in the drying process. This finding is explained by a soil-plant hydraulic model that assumes that stomata start to close when the soil hydraulic conductance of the soil-plant continuum starts to decline. Those findings stress the importance of belowground hydraulic properties on stomatal regulation and thereby drought responsiveness.
- 27 March 2022
- Published by Copernicus GmbH
Open AccessResponsiveness of maize to soil drying is related to a decrease in belowground hydraulic conductivity
Limited water supply is one of the largest impediments to food production worldwide in the light of climate change and increasing food demand. Stomatal regulation allows plants to promptly react to water stress and regulate water use. Although the coordination between stomatal closure and aboveground hydraulics has extensively been studied, our understanding of the impact of belowground hydraulics on stomatal regulation remains, as yet, incomplete. The overall objective of this study was to investigate the impact of belowground hydraulic conductivity as affected by differences in expressions of root and rhizosphere traits on the water use regulation of different maize genotypes.We have utilized a novel phenotyping facility to investigate the response of a selection of 48 maize (Zea mays L.) genotypes exhibiting different root and rhizosphere traits to soil drying. We measured the relation between leaf water potential, soil water potential, soil water content and transpiration rate, as well as root and rhizosphere traits (e.g. root length, rhizosheath mass) between genotypes. Our hypothesis is that stomatal response to soil drying is related to a loss in soil hydraulic conductivity and that key root and rhizosphere hydraulic traits affect such relation.We found that the genotypes differed in their responsiveness to drought and that such differences were related to belowground hydraulic traits. The critical soil water content at which plants started to decrease transpiration was related to a combination of plant- and rhizosphere traits (namely plant hydraulic conductance, maximum transpiration rate, root length and rhizosheath mass). Genotypes with a higher maximum transpiration rate and a higher plant hydraulic conductance and a smaller root system closed stomata in wetter soil conditions, meaning earlier in the drying process. This finding is explained by a soil-plant hydraulic model that assumes that stomata start to close when the soil hydraulic conductance of the soil-plant continuum starts to decline. Those findings stress the importance of belowground hydraulic properties on stomatal regulation and thereby drought responsiveness.