Does the root to shoot ratio show a hormetic response to stress? An ecological and environmental perspective

Root/shoot(R/S)ratio is an important index for assessing plant health,and has received increased attention in the last decades as a sensitive indicator of plant stress induced by chemical or physical agents.The R/S ratio has been discussed in the context of ecological theory and its potential importance in ecological succession,where species follow different strategies for above-ground growth for light or below-ground competition for water and nutrients.We present evidence showing the R/S ratio follows a biphasic dose–response relationship under stress,typical of hormesis.The R/S ratio in response to stress has been widely compared among species and ecological succession classes.It is constrained by a variety of factors such as ontogeny.Furthermore,the current literature lacks dose-response studies incorporating the full dose–response continuum,hence limiting scientific understanding and possible valuable application.The data presented provide an important perspective for new-generation studies that can advance current ecological understanding and improve carbon storage estimates by R/S ratio considerations.Hormetic response of the R/S ratio can have an important role in forestry for producing seedlings with desired characteristics to achieve maximum health/productivity and resilience under plantation conditions.

Water Deficit Stress Effects on Corn (<i>Zea mays</i>, L.) Root:Shoot Ratio

A study was conducted at Akron, CO, USA, on a Weld silt loam in 2004 to quantify the effects of water deficit stress on corn (Zea mays, L.) root and shoot biomass. Corn plants were grown under a range of soil bulk density and water conditions caused by previous tillage, crop rotation, and irrigation management. Water deficit stress (Dstress) was quantified by the number of days when the water content in the surface 0.3 m deviated from the water content range determined by the Least Limiting Water Range (LLWR). Root and shoot samples were collected at the V6, V12, and R1 growth stages. There was no significant correlation between Dstress and shoot or root biomass at the V6 growth stage. At the V12 and R1 growth stages, there were negative, linear correlations among Dstress and both root biomass and shoot biomass. The proportional decrease of shoot biomass was greater than the proportional decrease in root biomass, leading to an increase in the root:shoot ratio as water deficit stress increased at all growth stages. Determining restrictive soil conditions using the LLWR may be useful for evaluating improvement or degradation of the soil physical environment caused by soil management.

Establishment of highly efficient plant regeneration of Paeonia ostii‘Fengdan’through optimization of callus,adventitious shoot,and rooting induction

Tree peony is a famous ornamental plant,while the low propagation rate is the main hurdles hindering the industry development.Till now,the highly efficient regeneration system for tree peony is not established.In this study,using Paeonia ostii’Fengdan’mature embryos,the effects of variations in inoculation method,initiating culture,adventitious shoot induction,rooting media,plant growth regulators(PGRs),and a nonconventional PGR(plant extracts)on regeneration from explants were evaluated.In embryo cultures,embryonic callus induction rate of 1/4 embryos was the highest among those of embryos with other three technical treatments(whole embryos,1/2 embryos,and pieces of embryos).The woody plant medium(WPM)containing 1.0 mg·L^(-1)6-BA,0.5 mg·L^(-1)GA3,30.0 g·L^(-1)sucrose,and 3.0 g·L^(-1)phytagel significantly improved shoot induction and multiplication.3.0 mg·L^(-1)plant extracts promoted hypocotyl germination,rooting,and root growth,in direct embryo culture,and a combination of 3.0 mg·L^(-1)plant extracts+2.0 mg·L^(-1)IBA+1.5 mg·L^(-1)IAA produced optimal rooting induction rate for multiple shoots in direct embryo culture and indirect somatic embryogenesis.For the three in vitro micropropagation methods,the highest shoot proliferation coefficient(5.4±0.2)was obtained with indirect somatic embryogenesis.Fortunately,the propagation ability of shoots remains high,even when culture propagation was continued for more than two years.Thus,a reliable system for plant regeneration from mature embryos derived from P.ostii’Fengdan’callus and two direct embryo culture systems have been established.The novel regeneration system could facilitate uniform seedling production.

Regulation of phytohormones on root primordium initiation and adventitious root formation in the etiolated shoots of Paeonia suffruticosa'Yinfen Jinlin'

Tree peony is well known and sought after for its large, colorful flowers. Its propagation is via vegetative methods. Mech- nisms of the adventitious rooting and the regulation of rooting processes are the principles and techniques of plant propagation and improvement. Microstructures and fluctuations of phytohormones in the adventitious rooting were studied with the etiolated soft- wood shoots of Paeonia suffkuticosa ＇Yinfen Jinlin＇. There are no pre-primordia in the shoots of the cultivar. Adventitious roots are produced in five stages： shoot selection, primordium initiation, primordium growth, conducting tissue differentiation and root protru- sion. Primordia initiated in the cortex. The contents of the endogenous hormones, IAA, ABA and GA, were 5.842, 0.873 and 1.043 nmol·g^-1 FW on the bases of shoots, respectively. CTKs which included isopentenyl adenine （iPA）, zeatin riboside （ZR） and dihy- drozeatin riboside （DHZR） were 0.949, 0.695 and 2.034 nmol·g^-1 FW, respectively. DHZR is active among CTKs. The ratio of IAA to GA, CTK and ABA clearly increased at the stage of primordium initiation, while they showed low levels at the stages of primor- dium growth. The ratios were restored at the shoot levels at the stage of root protrusion. IBA provoked primordia initiation in the cortex, the vascular cambium, the pith and even in the callus induced on the base of shoots. ]AA levels in the treated shoots increased gradually to its highest level （three times of control） at the stage of conducting tissue differentiation. The ratios of IAA to GA, CTK and ABA clearly decreased at the stage of primordium initiation. The ratio of IAA to ABA is regulated at 10：1.

Effects of Free-air CO2 Enrichment on Root Characteristics and C:N Ratio of Rice at the Heading Stage

A hydroponics experiment was conducted to investigate the rice root growth in FACE （free-air carbon dioxide enrichment）. The root biomass, root volume, ratio of root/shoot, number of adventitious roots and root diameter significantly increased under FACE conditions, while the CO2 enrichment decreased the N concentration in rice roots without any change in the C content, leading to an increase in root C：N ratio. Moreover, the elevated CO2 resulted in a remarkable decrease of root activity, expressed as per unit root dry weight, which might be responsible for decreased N concentration in roots.

The Equilibrium and Growth Stability of Winter Wheat Root and Shoot Under Different Soil Water Conditions

The equilibrium between root, shoot and growth stability under different soil water conditions were investigated in a tube experiment of winter wheat. The water supplying treatments included： sufficient irrigation at whole growth phase, moderate deficiency irrigation at whole growth phase, serious deficiency irrigation at whole growth phase, sufficient irrigation at jointing stage, tillering stage, flowering stage, and fillering respectively, after moderate and serious water deficit during their previous growth stage. Root and shoot biomass were measured. On the basis of the cooperative root-shoot interactions model, the equilibrium and growth stability were studied on the strength of the kinetics system theory. There was only one varying equilibrium point between the root and shoot over the life time of the winter wheat plant. Water stress prolonged the duration of stable growth, the more serious the water deficit, the longer the period of stable growth. The duration of stable growth was shortened and that of unstable growth was prolonged after water recovery. The growth behavior of the plants exposed to moderate water deficit shifted from stable to unstable until the end of the growth, after rewatering at flowering. In the life-time of the crop, the root and shoot had been adjusting themselves in structure and function so as to maintain an equilibrium, but could not achieve the equilibrium state for long. They were always in an unbalanced state from the beginning to the end of growth. This was the essence of root-shoot equilibrium. Water stress inhibited the function of root and shoot, reduced root shoot interactions, and as a result, the plant growth gradually tended to stabilize. Rewatering enhanced root shoot interactions, prolonged duration of instable growth. Rewatering at flowering could upset the inherent relativity during the long time of stable growth from flowering to filling stage, thus leading to unstable growth and enhanced dry matter accumulating rate in the whole plant.

Effects of Soil Moisture on Dynamic Distribution of Dry Matter Between Winter Wheat Root and Shoot

The dynamic relationship of dry matter accumulation and distribution between winter wheat root and shoot was studied under different soil water conditions. The dry matter accumulation in root was greatly influenced by water stress, so as to the final root weight of the treatment with 40% field moisture capacity(FMC) was less than 1/4 of that of the treatment with 80% FMC on average. Water stress during the 3-leaf stage to the tillering stage had the greatest influence on root, and the influence of water stress during the jointing stage to the booting stage on shoot was greater than root. However, water stress during the tillering stage to the booting stage had a balanced effect on root and shoot, and the proportion of dry matter that distributed to root and shoot was almost the same after rewatering. Water recovery during the jointing stage to booting stage could promote R/S, but the increasing degree was related to the duration of water limitation. Soil water condition had the lowest effect on R/S during the flowering stage to the filling stage and the maximal effect on R/S during the jointing stage to the heading stage, R/S of 40% FMC treatment was 20.93 and 126.09% higher than that of 60% FMC and 80% FMC treatments respectively at this period.

Differential responses of root growth to nutrition with different ammonium/nitrate ratios involve auxin distribution in two tobacco cultivars

Nitrogen(N), the major forms of which are nitrate(NO3–) and ammonium(NH4+), plays an important role in plant growth and mediation of root development. However, the role of auxin in root growth in response to different NH4+/NO3– ratios remains unclear. Two tobacco cultivars(Nicotiana tabacum L.) were adopted in this study, which displayed variant growth features under the situations with sole NO3– nutrition ratio(NH4+/NO3– ratio: 0/100), low NO3– nutrition ratio(NH4+/NO3– ratio: 97/3), and optimal NH4+/NO3– ratio(50/50). We investigated the effects of the different NH4+/NO3– ratios on the formation and elongation of lateral roots(LRs), auxin concentration, DR5::GUS expression, 3 H-labeled indole acetic acid([3 H]IAA) transport, and the expression of six PIN genes in tobacco roots. We also examined the effects of exogenous auxin and a transport inhibitor on LRs growth. The results are shown as follows, compared to optimal N nutrition conditions, the biomass and nitrogen(N) accumulation were largely reduced by sole and low NO3– nutrition treatment in NC89, but no difference was observed in Zhongyan 100. In most cases, sole and low NO3– nutrition impaired the elongation and formation of firstorder lateral roots(1° LRs), only in NC89, thus reducing the root growth. IAA concentration and DR5::GUS expression levels decreased in roots when NC89 was subjected to sole and low NO3– nutrition media, suggesting that different NH4+/NO3– ratios affect the transport of auxin from leaves to roots. Results were similar following exogenous NAA application to low NO3– nutrition treated seedlings. Based on direct [3 H]IAA transport measurement, the transport of polar auxin from shoots to roots decreased due to low NO3– nutrition. PIN4 expression levels were markedly decreased in roots of NC89by sole and low NO3– nutrition, while they were unaffected in Zhongyan 100 roots. Overall, our findings suggest that LRs formation in tobacco seedlings is regulated by NH4+/NO3– ratios via modifying polar transport of auxin.

Direct Regeneration of Plants Derived from in vitro Cultured Shoot Tips and Leaves of Poplar （Populus×euramericana ＇Neva＇）

The purpose of the present study was to establish a regeneration procedure for Populus × euramericana ＇Neva＇ by using in vitro shoots tips and leaves. For sterilization, 0.1% （w/v） mercuric chloride （HgCl2） solution for 8 to 10 min was the optimal treatment for this poplar cultivation. The effects of benzyladenine （BA） and α-naphthaleneacetic acid （NAA） added to Murashige and Skoog （MS） medium were tested on organogenesis. The highest regeneration rate and numbers of shoots/explant from shoot tips （96.7%, 9.8） and leaves （90.0%, 8.7） were obtained on the half-strength MS medium supplemented with 0.5 mg/L BA and 0.1 mg/L NAA. The optimal medium for in vitro rooting of shoots was on a half-strength MS medium containing 1 mg/L indolebutyric acid （IBA） with the highest rooting frequency （93.3%） and numbers of roots/explant （8.2）. For acclimatization, in vitro rooted plantlets were transferred to plastic cups containing vermiculite and peat （1： 1）. After acclimatization, transplanted plantlets grew well in a shade house. Therefore, we believe that this efficient plant regeneration protocol especially by leaf explants is very important for in vitro clonal propagation of Populus×euramericana ＇Neva＇.

Root-shoot relationships of Fraxinus mandchurica

In this paper, root-shoot relationships of seedlings of Manchurican ash (Fraxinus mandchurica) in pots is studied in green house. The results show that roots and shoots have the co-ordination of growth and roots and shoots of ash have the same growth dynamics. There are very close relationships between toall root mass and total shoot mass, between fine root mass and leaf mass in this experiments. At the end growing season, ash root-shoot mass ratio and fine root-leaf mass ratio are 1 .85 and 2.01 respectively. These ratios are difference during growth season. There are larger roots than shoots on seedling stage. Low nitrogen and phosphorus produce higher root-shoot ratio, and high nutrient concentrations decrease the ratios.

Variations in shoot and root growth of three provenances of Faidherbia albida in clay and sand soil

The variation in shoot and root seedling growth traits was compared among three Faidherbia albida （Del）. A. Chev. provenances originating from Kenya （Taveta）, Malawi （Bolero） and Ghana （Bolgatanga）; representing east, south and west Africa, respectively. Bulked seeds from three provenances were grown in two soil types （clay and sand） at the World Agroforestry Centre nursery, Nairobi. Seedlings grown in clay soil had high shoot growth and shoot to root ratios but shorter root length, lower number of nodules and specific root length compared to seedlings in sand soil. Shoot to root ratios reduced with time and this was more pronounced in sand soil. Overall, the Bolero and Taveta provenances had higher growth than the Bolgatanga provenance. This was probably because plants grown in nutrient-rich clay soil invest more in shoot growth as compared to root growth. Longer root growth in sand soil shows a better adaptive mechanism that increases competitive ability and survival in nutrient-poor systems. Variations observed among the seedling variables such us shoot and root growth could be used for early selection for reforestation and agroforestry in predominantly clay and sand soil areas.

Phytohormones Accumulation and Distribution in Shoots and Roots of Haploid,Diploid and Tetraploid Barley Seedlings Derived from Microspore Culture

Phytohormones play important roles in plant growth and development,and polyploids are thought to be an important method for plant breeding.However,the relationship between ploidy and phytohormone is still unclear.In this study,barley at three ploidy levels were produced by microspore culture.Therefore,we further analyzed the phytohormone content in the shoots and roots of the three kinds of barley materials to study the effect of ploidy on phytohormones accumulation and distribution.The results showed that Abscisic acid(ABA),gibberellin(GA),jasmonic acid(JA),auxin(IAA),salicylic acid(SA)and cytokinin(CTK)were successfully determined in shoots and roots using LC-MS(liquid chromatography mass spectrometry).By comparing the shoots of the haploid and diploid plants,it was found that the distribution trend of the six phytohormones was consistent,and another consistent trend was found in the roots of the diploid and tetraploid plants.In addition,we further analyzed the shoot/root ratio of the different phytohormones to identify the potential differences for haploid,diploid and tetraploid.Here,the relationship between ploidy and phytohormone we provided would provide new insights into understanding the new phenotypes that occur in polyploid species.

Root to crown ratio of permanent teeth of people living in Shanghai

The aim of this study is to assess if there is a significant difference between crown to root ratio of male and female permanent teeth in Chinese people in Shanghai.Methods: For this purpose one thousand four hundred (700,male and 700,female) extracted teeth were collected between age 25-35 and the crown length (CL) and root length (RL) of permanent teeth were measured by a precise caliper.Statistical analyses were performed with the SPSS 17 software by Student's t-test method.Results: The mean crown length of permanent teeth of male was found to be larger than that of female except for maxillary central incisor and the mean root length of male was found to be comparatively larger than that of female.Conclusion: There was a significant difference in root to crown ratio (RL/CL) between male maxillary teeth and mandibular teeth and female ones.The RL/CL ratio of male tends to be larger than those of female in Shanghai population.

Effect of Shoot-Fruit Ratio on Grape Yield,Quality and Storage Nutrient

With the grape variety ＂Summer Black＂ as the test material, the effects of different shoot-fruit ratios on grape yield, fruit quality and grapevine storage nutrient were investigated for 2 consecutive years, so as to determine the optimized load for Summer Black grapes in Shandong. The results showed that the grapes with shoot-fruit ratio of 1∶ 1 had the highest soluble solid content, vitamin C content and grapevine storage nutrient. Based on the comprehensive analysis, the shoot-fruit ratio of 1∶ 1 was the optimized load for Summer Black grapes in Tai an of Shandong Province, and the yield was 1 179.37-1 288.63 kg per 667 m^2.

Root overlap and allocation of above- and belowground growth of European beech in pure and mixed stands of Douglas fir and Norway spruce

Site conditions and species identity have a combined effect on fine root growth of trees in pure and mixed stands.However,mechanisms that may contribute to this effect are rarely studied,even though they are essential to assess the potential of species to cope with climate change.This study examined fine root overlap and the linkage between fine root and stem growth of European beech(Fagus sylvatica)growing in pure and mixed stands with Douglas fir(Pseudotsuga menziesii)or Norway spruce(Picea abies)at two different study sites in northwestern Germany.The study sites represented substantially different soil and climate conditions.At each site,three stands,and at each stand,three pairs of trees were studied.In the pure beech stand,the pairs consisted of two beech trees,while in the mixed stands each pair was composed of a beech tree and a conifer.Between each pair,three evenly spaced soil cores were taken monthly throughout the growing season.In the pure beech stands,microsatellite markers were used to assign the fine roots to individual trees.Changes in stem diameter of beech were quantified and then upscaled to aboveground wood productivity with automatic high-resolution circumference dendrometers.We found that fine root overlap between neighboring trees varied independently of the distance between the paired trees or the stand types(pure versus mixed stands),indicating that there was no territorial competition.Aboveground wood productivity(wood NPP)and fine root productivity(root NPP)showed similar unimodal seasonal patterns,peaking in June.However,this pattern was more distinct for root NPP,and root NPP started earlier and lasted longer than wood NPP.The influence of site conditions on the variation in wood and root NPP of beech was stronger than that of stand type.Wood NPP was,as expected,higher at the richer site than at the poorer site.In contrast,root NPP was higher at the poorer than at the richer site.We concluded that beech can respond to limited resources not only above-but also belowground and that the negative relationship between above-and belowground growth across the study sites suggests an‘optimal partitioning’of growth under stress.

Lateral root elongation in maize is related to auxin synthesis and transportation mediated by N metabolism under a mixed NO_(3)^(–) and NH_(4)^(+) supply

A mixed nitrate (NO_(3)^(–)) and ammonium (NH_(4)^(+)) supply can promote root growth in maize (Zea mays),however,the changes in root morphology and the related physiological mechanism under different N forms are still unclear.Here,maize seedlings were grown hydroponically with three N supplied in three different forms (NO_(3)^(–)only,75/25 NO_(3)^(–)/NH_(4)^(+)and NH_(4)^(+)only).Compared with sole NO_(3)^(–)or NH_(4)^(+),the mixed N supply increased the total root length of maize but did not affect the number of axial roots.The main reason was the increased total lateral root length,while the average lateral root (LR) length in each axle was only slightly increased.In addition,the average LR density of 2nd whorl crown root under mixed N was also increased.Compared with sole nitrate,mixed N could improve the N metabolism of roots (such as the N influx rate,nitrate reductase (NR) and glutamine synthase (GS)enzyme activities and total amino content of the roots).Experiments with exogenously added NR and GS inhibitors suggested that the increase in the average LR length under mixed N was related to the process of N assimilation,and whether the NR mediated NO synthesis participates in this process needs further exploration.Meanwhile,an investigation of the changes in root-shoot ratio and carbon (C) concentration showed that C transportation from shoots to roots may not be the key factor in mediating lateral root elongation,and the changes in the sugar concentration in roots further proved this conclusion.Furthermore,the synthesis and transportation of auxin in axial roots may play a key role in lateral root elongation,in which the expression of ZmPIN1B and ZmPIN9 may be involved in this pathway.This study preliminarily clarified the changes in root morphology and explored the possible physiological mechanism under a mixed N supply in maize,which may provide some theoretical basis for the cultivation of crop varieties with high N efficiency.

Establishment of Somatic Cell Clones in Thesium chinense Turcz and Its in vitro Rooting Technique

[Objective] The aim of this study is to establish the rapid micro-propagation system in Thesium chinense Turcz.[Method]With stem fragments of wild Thesium chinense Turcz as explants,different culture media were designed to conduct induction culture,strengthening plantlet culture and in vitro rooting.[Result]The optimum medium for inducing clustered shoots was determined to be MS medium appended with 1.5 mg/L 6-BA,0.01 mg/L NAA and 0.3 mg/L 2,4-D;in addition,60 mg/kg ABT was suitable for rooting,by which the percentage of rooted plantlets reached 76.6%.[Conclusion]This study simplified the procedures of tissue culture in Thesium chinense Turcz and enhanced the proliferation rate,providing basis for artificial cultivation and resource protection of Thesium chinense Turcz.

多花胡枝子超嫩梢扦插繁殖及生根机理研究

探究植物生长调节剂对多花胡枝子超嫩梢扦插生根效果与内源激素含量变化的影响,揭示多花胡枝子超嫩梢扦插繁殖的生根机理。以多花胡枝子顶部嫩梢为材料,测定植物生长调节剂种类、浓度对多花胡枝子扦插生根率、生长及内源激素含量的影响。结果表明,多花胡枝子超嫩梢扦插生根类型为愈伤组织生根;生根过程经历愈伤组织阶段、不定根形成阶段和不定根伸长阶段;T5处理的插穗表现出最佳效果,生根率为100%。插穗内源激素与生根密切相关,与CK相比,处理组IAA与GA的含量较高,且IAA/ABA与IAA/ZR的含量比值较高,ABA与ZR的含量较低,经过植物生长调节剂处理后,有利于多花胡枝子超嫩梢扦插生根。研究获得了多花胡枝子超嫩梢扦插生根的最适宜处理,T5处理下有利于一级根生长,促进扦插苗的生长,提高IAA含量及其与ABA、ZR的比值,有利于多花胡枝子超嫩梢扦插生根。

A Comparative Study of Morphological Responses of Seedling Roots to Drying Soil in 20 Species from Different Habitats

Rooting depth and root and shoot biomass were measured for seedlings of 20 species in both watered and unwatered sand columns. The species were from habitats of widely varying moisture status, ranging from marsh to desert. Moisture status of the species' habitats was quantified as Ellenberg moisture number. Seedlings were allowed to grow in moist sand for 21 days and were then exposed to the treatments (watered and unwatered) for a further 21 days. Rooting depth of control plants was not correlated with Ellenberg number. Root depth of plants from dry habitats tended to increase in drying sand, while roots of plants from wet habitats decreased in depth. Plasticity of rooting depth (depth in unwatered / depth in watered sand) was significantly correlated with Ellenberg number ( r 2 = 0.56). Plasticity of shoot/root ratio was also correlated with Ellenberg number, but the relationship was weaker than for rooting depth plasticity. Species that showed the greatest plasticity in rooting depth also showed the greatest ability to sustain shoot growth in unwatered sand. There was some evidence that growth of plants from very dry habitats was reduced in the watered treatment. Results of this study suggest that a major, although not the only, adaptation of plants of dry habitats is the ability of their seedlings to exploit deeply buried water resources.

Study on the Influencing Factors and the Root Mechanism in Cutting Process of Prunus humilis(Bge). Sok

A procedure for cutting Prunus humilis（Bge）. Sok was comprehensively studied in this paper. It was found that the key factors involved in this procession were medium, rooting accelerator, concentration of rooting accelerator and type of shoot. The results showed that send was used as mediums; Treatment with 1 000 mg/L AST rooting powder No. 2 and semi-woody shoots were the optimal materials for cutting, and the rooting rate reached 88.1%. Anatomical study on rooting of Prunus hum#is（Bge）. Sok cutting has been carded out by the paraffin section method. The observation result shows that the adventitious root primordium of Prunus humili$（Bge）. Sok cutting belongs to the type of induced root primordium. The adventitiousroot primordium originates from the cross region of vascular cambium and pith rays.