Nutrient-hormone relations:Driving root plasticity in plants

Optimal plant development requires root uptake of 14 essential mineral elements from the soil.Since the bioavailability of these nutrients underlies large variation in space and time,plants must dynamically adjust their root architecture to optimize nutrient access and acquisition.The information on external nutrient availability and whole-plant demand is translated into cellular signals that often involve phytohormones as intermediates to trigger a systemic or locally restricted developmental response.Timing and extent of such local root responses depend on the overall nutritional status of the plant that is transmitted from shoots to roots in the form of phytohormones or other systemic long-distance signals.The integration of these systemic and local signals then determines cell division or elongation rates in primary and lateral roots,the initiation,emergence,or elongation of lateral roots,as well as the formation of root hairs.Here,we review the cascades of nutrient-related sensing and signaling events that involve hormones and highlight nutrient-hormone relations that coordinate root developmental plasticity in plants.

Use of genotype-environment interactions to elucidate the pattern of maize root plasticity to nitrogen deficiency

Maize（Zea mays L.） root morphology exhibits a high degree of phenotypic plasticity to nitrogen（N） de ficiency,but the underlying genetic architecture remains to be investigated Using an advanced BC_4F_3 population,we investigated the root growth plasticity under two contrasted N levels and identi fied the quantitative trait loci（QTLs） with QTL-environment（Q×E）interaction effects. Principal components analysis（PCA） on changes of root traits to N de ficiency（D LN-HN） showed that root length and biomass contributed for 45.8% in the same magnitude and direction on the first PC,while root traits scattered highly on PC_2 and PC_3. Hierarchical cluster analysis on traits for D LN-HN further assigned the BC_4F_3 lines into six groups,in which the special phenotypic responses to N de ficiency was presented These results revealed the complicated root plasticity of maize in response to N de ficiency that can be caused by genotype environment（G×E） interactions. Furthermore,QTL mapping using a multi-environment analysis identi fied 35 QTLs for root traits. Nine of these QTLs exhibited signi ficant Q×E interaction effects. Taken together,our findings contribute to understanding the phenotypic and genotypic pattern of root plasticity to N de ficiency,which will be useful for developing maize tolerance cultivars to N de ficiency.

Responses of root growth of Alhagi sparsifolia Shap. (Fabaceae) to different simulated groundwater depths in the southern fringe of the Taklimakan Desert, China

Alhagi sparsifolia Shap. （Fabaceae） is a spiny, perennial herb. The species grows in the salinized, arid regions in North China. This study investigated the response characteristics of the root growth and the dis- tribution of one-year-old A. sparsifolia seedlings to different groundwater depths in controlled plots. The eco- logical adaptability of the root systems of A. sparsifolia seedlings was examined using the artificial digging method. Results showed that： （1） A. sparsifolia seedlings adapted to an increase in groundwater depth mainly through increasing the penetration depth and growth rate of vertical roots. The vertical roots grew rapidly when soil moisture content reached 3%-9%, but slowly when soil moisture content was 13%-20%. The vertical roots stopped growing when soil moisture content reached 30% （the critical soil moisture point）. （2） The morphological plasticity of roots is an important strategy used by A. sparsifolia seedlings to obtain water and adapt to dry soil conditions. When the groundwater table was shallow, horizontal roots quickly expanded and tillering increased in order to compete for light resources, whereas when the groundwater table was deeper, vertical roots developed quickly to exploit space in the deeper soil layers. （3） The decrease in groundwater depth was probably respon- sible for the root distribution in the shallow soil layers. Root biomass and surface area both decreased with soil depth. One strategy of A. sparsifolia seedlings in dealing with the increase in groundwater depth is to increase root biomass in the deep soil layers. The relationship between the root growth/distribution of A. sparsifolia and the depth of groundwater table can be used as guidance for harvesting A. sparsifolia biomass and managing water resources for forage grasses. It is also of ecological significance as it reveals how desert plants adapt to arid environments.

Plasticity of wheat seedling responses to K^(+) deficiency highlighted by integrated phenotyping of roots and root hairs over the whole root system

The availability in the soil of potassium(K^(+)),a poorly mobile macronutrient required in large quantities for plant growth,is generally suboptimal for crop production in the absence of fertilization,making improvement of the ability of crops to adapt to K^(+)deficiency stress a major issue.Increasing the uptake capacity of the root system is among the main strategies to achieve this goal.Here,we report an integrative approach to examine the effect of K^(+)deficiency on the development of young plant entire root system,including root hairs which are known to provide a significant contribution to the uptake of poorly mobile nutrients such as K^(+),in two genetically distant wheat varieties.A rhizobox-type methodology was developed to obtain highly-resolved images of root and root hairs,allowing to describe global root and root hair traits over the whole root system via image analysis procedures.The two wheat varieties responded differently to the K^(+)shortage:Escandia,a wheat ancestor,reduced shoot biomass in condition of K^(+)shortage and substantially increased the surface area of its root system,specifically by increasing the total root hair area.Oued Zenati,a landrace,conversely appeared unresponsive to the K^(+)shortage but was shown to constitutively express,independently of the external K^(+)availability,favorable traits to cope with reduced K^(+)availability,among which a high total root hair area.Thus,valuable information on root system adaptation to K^(+)deficiency was provided by global analyses including root hairs,which should also be relevant for other nutrient stresses.

Drought adaptability of phreatophytes:insight from vertical root distribution in drylands of China

Aims The vertical distribution of plant roots is a comprehensive result of plant adaptation to the environment.Limited knowledge on fine vertical root distributions and complex interactions between roots and environmental variables hinders our ability to reliably predict climatic impacts on vegetation dynamics.This study attempts to understand the drought adaptability of plants in arid areas from the perspective of the relationship between vertical root distribution and surroundings.Methods By analyzing root profiles compiled from published studies,the root vertical profiles of two typical phreatophytes,Tamarix ramosissima and Populus euphratica,and their relationships with environmental factors were investigated.A conceptual model was adopted to link the parameter distribution frequency with plant drought adaptability.Important Findings The strong hydrotropism(groundwater-dependent)and flexible water-use strategy of T.ramosissima and P.euphratica help both species survive in hyperarid climates.The differences in the developmental environments between T.ramosissima and P.euphratica can be explained well by the different distribution characteristics of root profiles.That is,higher root plasticity helps T.ramosissima develop a more efficient water-use strategy and therefore survive in more diverse climatic and soil conditions than P.euphratica.We conclude that the higher variation in root profile characteristics of phreatophytes can have greater root adaptability to the surroundings and thus wider hydrological niches and stronger ecological resilience.The inadequacy of models in describing root plasticity limits the accuracy of predicting the future response of vegetation to climate change,which calls for developing process-based dynamic root schemes in Earth system models.

Evaluation of high yielding soybean germplasm under water limitation

Limited information is available for soybean root traits and their plasticity under drought stress. To date, no studies have focused on examining diverse soybean germ- plasm for regulation of shoot and root response under water limited conditions across varying soil types. In this study, 17 genetically diverse soybean germplasm lines were selected to study root response to water limited conditions in clay （trial 1） and sandy soil （trial 2） in two target environments. Physiological data on shoot traits was measured at multiple crop stages ranging from early vegetative to pod filling. The phenotypic root traits, and biomass accumulation data are collected at pod filling stage. In trial 1, the number of lateral roots and forks were positively correlated with plot yield under water limitation and in trial 2, lateral root thickness was positively correlated with the hill plot yield. Plant Introduction （PI） 578477A and 088444 were found to have higher later root number and forks in clay soil with higher yield under water limitation, in sandy soil, P1458o2o was found to have a thicker lateral root system and higher yield under water limitation. The genotypes identified in this study could be used to enhance drought tolerance of elite soybean cultivars through improved root traits specific to target environments.