Chloropicrin Soil Fumigation Reduces <Emphasis Type="Italic">Spongospora subterranea</Emphasis> Soil Inoculum Levels but Does Not Control Powdery Scab Disease on Roots and Tubers of Potato

The effect of chloropicrin fumigation on the soil populations of Spongospora subterranea and the development of powdery scab, formation of root galls and tuber yield was investigated in seven field trials conducted in Minnesota and North Dakota. Sixteen potato cultivars, with different levels of susceptibility to disease on roots and tubers, were planted in plots treated with chloropicrin at rates ranging from zero to 201.8 kg a.i. ha^?1. The amount of S. subterranea DNA in soil was determined using qPCR. Bioassays were conducted to further assess the effect of chloropicrin fumigation on root colonization by S. subterranea in two potato cultivars with contrasting disease susceptibility. In the field, chloropicrin applied at rates between 70.1 to 201.8 kg a.i. ha^?1 significantly decreased S. subterranea initial inoculum in soil but increased the amount of disease observed on roots and tubers of susceptible cultivars. The effect of increasing disease was confirmed in controlled conditions experiments. Although the amount of S. subterranea DNA in roots of bioassay plants increased with increasing chloropicrin rates, it remained similar among potato cultivars. Chloropicrin fumigation significantly increased tuber yield which in cultivars such as Shepody and Umatilla Russet were associated with the amount root galls (r = 0.30; P < 0.03). Results of these studies contradict earlier reports on the use of chloropicrin fumigation for the control of powdery scab. Factors other than inoculum level, such as environmental conditions that affect inoculum efficiency and host susceptibility, may be significant contributors to the development of powdery scab and root gall formation.     

Effects of Different Soil Treatments on the Development of <Emphasis Type="Italic">Spongospora subterranea</Emphasis> f. sp. <Emphasis Type="Italic">subterranea</Emphasis> in Potato Roots and Tubers in the Greenhouse

Powdery scab caused by Spongospora subterranea f. sp. subterranea (Sss) causes extensive losses in potato production systems globally. Two pot experiments were established in the greenhouse in summer 2013 and winter 2014 to evaluate the effectiveness of different soil chemicals, fumigant, amendments and biological control agents (BCAs) against Sss in the rhizospheric soil, potato roots and tubers. The study used visual assessment methods to assess the effect of treatments on root galling and zoosporangia production, and qPCR to measure Sss concentration in the soil and in the potato roots and tubers. All six soil treatments, namely metam sodium, fluazinam, ZincMax, calcium cyanamide, Biocult and a combination of Bacillus subtilis and Trichoderma asperellum recorded significantly (P < 0.05) lower numbers of zoosporangia in the roots compared to the untreated control. The same effect was observed on the concentration of Sss DNA in the roots at tuber initiation. A more diverse picture was obtained when root gall scores at tuber initiation and Sss DNA in the rhizospheric soil at tuber initiation and harvesting were compared. Significant differences (P < 0.05) were also noted in disease severity, disease incidence, and tuber yield between metam sodium, fluazinam, ZincMax, calcium cyanamide and the untreated control. Calcium cyanamide gave the highest tuber yield. The study demonstrated the potential of soil treatments such as metam sodium, fluazinam, ZincMax and calcium cyanamide in managing Sss in potatoes by reducing the pathogen both in the rhizospheric soil and the roots of the potato plant.     

Effect of cultivar maturity period on the growth and yield of potato plants grown from microtubers and conventional seed tubers

Potato plants of early cultivars grown from microtubers have been reported to have a much lower growth vigor and produce lower yields than microtubers of late cultivars. This study intended to clarify the field performance of plants grown from directly planted microtubers of cultivars with different maturity periods, with a special attention to early cultivars. The experiments were conducted at Hokkaido University, Japan, over four years. Microtubers and conventional seed tubers of the early cultivar Kitaakari, late cultivars Konafubuki and Norin 1, and very late breeding line IWA-1 were planted, and the plant growth and tuber yields were analyzed. The microtuber plants of Kitaakari had a lower initial increase in leaf area index than conventional seed tuber plants, but at the maximum shoot growth had the same leaf area index. This pattern was also observed in the other cultivars. Tuber initiation and tuber bulking occurred on average five days later in microtuber plants than in conventional seed tuber plants of cultivar Kitaakari. At maximum shoot growth, microtuber plants had on average 65% of tuber dry weight of conventional seed tuber plants, with small variation among cultivars. Irrespective of maturity period, microtuber plants showed a higher tuber increase after maximum shoot growth, achieving around 86% of tuber dry weight of conventional seed tuber plants at harvest. From the results of this study we conclude that microtuber plants of early and late cultivars have a similar yield potential relative to conventional seed tuber plants, and microtubers of both early and late cultivars might be used as an alternative seed tuber source for potato production, if necessary.     

孕穗期渍水对不同小麦品种根系生长和籽粒产量的影响

为给长江中下游地区小麦高产耐渍品种选用提供参考,以小麦品种扬麦25、扬麦24、宁麦13和宁麦9号为材料,分析了孕穗期连续10 d渍水处理对小麦叶面积和光合速率、地上部干物质积累、籽粒产量及其结构和不同土层（0~100 cm）根系干重的影响。结果表明,与对照（正常水分）相比,渍水处理显著降低小麦籽粒产量,降幅12%~31%,穗粒数和千粒重的减少是减产的主要原因。相比其他品种,渍水后扬麦25产量降幅小,产量构成较协调,受渍水影响轻。渍水处理显著减少小麦开花期和成熟期不同土层根系干重,其中对下层根系的影响大于上层根系;渍水也显著降低乳熟期倒三叶面积、倒二叶和倒三叶净光合速率,明显抑制成熟期地上部各器官和花后光合物质积累。在对照条件下,扬麦25的根系干重在开花期与其他品种相近,成熟期的0~20和80~100 cm土层根系干重则较高;在渍水条件下,扬麦25能在花后维持较高的上层根系生物量,成熟期的0~60 cm土层根干重占总根干重比例较高。不同水分处理下,扬麦25均具有面积大的旗叶且高的净光合速率,花后维持较久的上三叶绿色面积和积累更多的光合产物。因此,小麦花后根系生物量保持稳定（尤其是表层根系）,绿叶面积较大且衰减慢（尤其是旗叶）,有助于增强小麦花后光合物质积累能力,促进籽粒灌浆和单穗高产稳产,这可作为高产耐渍品种的筛选依据。     

Drought-induced root plasticity of two upland NERICA varieties under conditions with contrasting soil depth characteristics

To identify differences in root plasticity patterns of two upland New Rice for Africa (NERICA) varieties, NERICA 1 and 4, in response to drought under conditions with contrasting soil profile characteristics, soil moisture gradients were imposed using a sloping bed system with depths ranging 30–65 cm and a line-source sprinkler system with a uniformly shallow soil layer of 20 cm depth. Varietal differences in shoot and root growths were identified only under moderate drought conditions, 11–18% v/v soil moisture content. Further, under moderate drought soil conditions where roots could penetrate into the deep soil layer, deep root development was greater in NERICA 4 than in NERICA 1, which contributed to maintaining dry matter production. However, under soil conditions with underground impediment to deep root development, higher shoot dry weight was noted for NERICA 1 than for NERICA 4 at 11–18% v/v soil moisture content, which was attributed to increased lateral root development in the shallow soil layer in NERICA 1. Enhanced lateral root development in the 0–20-cm soil layer was identified in NERICA 1 even under soil conditions without an impediment to deep root development; however, this did not contribute to maintaining dry matter production in upland rice. Thus, we show different root developmental traits associated with drought avoidance in the two NERICA varieties, and that desirable root traits for upland rice cultivation vary depending on the target soil environment, such as the distribution of soil moisture and root penetration resistance.     

The growth of roots of perennial, Italian, hybrid and annual ryegrasses through a high-strength root medium

Abstract A test for root penetration through a wax disc of known hardness was developed and used with twenty‐five cultivars and twenty‐one breeding lines of perennial, Italian, hybrid and annual ryegrasses. After 42 days, the number of roots reaching the wax disc and the number penetrating the disc were counted. Root diameters were measured using image analysis. The proportion of root penetrations ranged from 0·06 to 0·39 across the cultivars and breeding lines, with substantial variation between replicates and cultivars. Diploid perennial cultivars had the thinnest roots, and there were no consistent differences in root diameter between perennial breeding lines, hybrid ryegrasses and Italian ryegrasses. Roots that penetrated the wax disc increased in diameter, by 0·60 on average, a few millimetres above the wax, and through the wax disc. Diameters below the wax disc were the same as those above the zone of impeded root growth. The increase was caused by an increase in size of the parenchyma cells of the root cortex. An experiment with partially and completely impeded root growth showed that impedance did not change root diameter of new roots, or the distribution of root mass between impeded and unimpeded halves of a root system for plants up to 10 weeks old.     

东北黑土区高产大豆R5期根系分布特征

利用钻土法及根系扫描分析系统研究不同产量类型大豆R5期根系在植株周围、株间及垄间土壤中的空间分布特征。结果表明：高产类型大豆根冠比较高，而且单位根长、根表面积的地上部干物质较高。根干重不仅在根系较集中的植株周围及株间0 ~ 30 cm的土层，而且在较深的土层（> 30 cm）中都有较多的分布；根长在植株周围的0 ~ 45 cm土层范围内表现出一定的优势。直径 < 1.0 mm的细根形成了根系的主要部分，根长差异的主要原因不是细根比例的大小，而是细根数量的多少。除株间0 ~ 15 cm土层外，高产大豆各土层的根/土体积比较高。     

Effects of Subsoiling to the Non-tilled Field of Wheat-Soybean Rotation on the Root System Development,Water Uptake,and Yield

Abstract

We introduced subsoiling to a field of wheat-soybean rotation where no-tillage practice had been conducted for five years and whose yield tended to decrease or stagnate. By subsoiling a half of each plot just before wheat sowing, treatments of tillage/no-tillage × subsoiling/no-subsoiling were established. Root distribution, shoot growth, water uptake and yield of both crops were examined to elucidate whether the subsoiling improves the productivity such as shoot biomass and yield through the modification of root system development, and how differ the effects of subsoiling between tilled and non-tilled fields. In wheat, roots were less concentrated in surface (0 ? 5 cm) layer in no-tillage, and distributed more in deep (20 ? 25 cm) layer of the soil. Deuterium labeled heavy water analysis revealed that the subsoiling enhanced water uptake from the deep soil layer in the no-tillage field. Both the no-tillage and subsoiling showed positive and significant effect on total biomass and yield. The effect of subsoiling must be related to water supply by deep roots in spring. In soybean no-tillage significantly increased the productivity, but subsoiling did not though distribution of the roots was modified by both practices. Soybean in non-tilled accumulated roots in the surface soil layer, but subsoiling did not significantly modify the root distribution especially in the deep soil layer. Water uptake trend and yield was thus not changed significantly by subsoiling. Subsoiling in the non-tilled field increased rooting depth and showed the possibility of braking yield stagnation in long-term no-tillage cultivation in wheat, but not in soybean.     

马铃薯抗旱资源的筛选与评价——营养体对块茎膨大速度的影响

摘要马铃薯地上部营养体是形成产量的重要因素之一,营养体与块茎膨是速度呈曲线正相关.营养体的形成随株龄的增长呈直线上升,品种不同则上升的程度也不同,中晚熟品种一般在8月下旬为最盛期,随后急剧下降.块茎形成膨大期因品种不同而异,膨大速度从现蕾期至成熟一直呈直线上升,品种不同则膨大速度各异.由此,根据营养体的量变数据可预测马铃薯块茎产量的高低、同时也可作为选育丰产型品种的一个重要依据.     

Assessing the tolerance toGlobodera pallida of resistant potato genotypes by means of field and pot tests

Summary The tolerance toGlobodera pallida of 15 potato genotypes with a high level of resistance toG. pallida was assessed by measuring their tuber yield in heavily infested fields and in nematicide treated strips in the same fields. Tolerance was also assessed in pots in the glasshouse by measuring the total biomass after 35 and 70 days growth in heavilyG. pallida infested soil and in uninfested soil. Large differences in tolerance between the genotypes were observed. High correlations were observed between tolerance assessments made in the field experiments and between field and pot experiments. No relationship was found between tolerance and maturity. Shoot and root dry weight were about equally affected by the nematode.     

Soil water stress and the growth and yield of potato plants grown from microtubers and conventional seed tubers

Tuber yields of potato plants grown from microtubers in fields are more variable than yields from conventional seed tubers (CT). One reason could be their higher susceptibility to water stress. This study clarified the effect of soil water stress from 1 month after emergence on the growth and yield of plants grown from conventional seed tubers and microtubers in fields. Microtubers (0.5–3 g) and conventional seed tubers (50 g) were grown in Hokkaido, Japan, over three field seasons. One month after emergence, poly-shelters were placed over the plots to prevent rainfall, and either irrigated (wet plot) or non-irrigated (dry plot) treatments were formed. At mid-flowering (about 50 days after emergence) leaf area index (LAI) in microtuber plants was decreased relatively more due to soil water stress than LAI in conventional seed tuber plants. However, at maximum shoot growth (about 80 days after emergence) both microtuber and conventional seed tuber plants had a similar relative decrease in LAI due to soil water stress. At mid-flowering and maximum shoot growth microtuber and conventional seed tuber plants had reduced stomatal conductance due to soil water stress, but the reduction in stomatal conductance was greater in conventional seed tuber plants than in microtuber plants. Microtuber and conventional seed tuber plants had similar root development at maximum shoot growth. Tuber production from mid-flowering until plant maturity was similarly affected by soil water stress in microtuber and conventional seed tuber plants. At harvest, plants affected by soil water stress had about 87% of the tuber dry weight of irrigated plants. We conclude, that the greater variation on tuber yield of microtuber plants cannot be attributed to soil water stress from 1 month after emergence.     

Light use efficiencies of potato cultivars with late blight (Phytophthora infestans)

Summary Potato cultivars of different maturity classes and levels of resistance toPhytophthora infestans were grown under several disease intensities in three field trials. Seasonal courses of ground cover by green foliage and final tuber yields were determined. Light use efficiencies (LUE) were calculated from regression analyses of yield on cumulative light interception. Late blight reduced tuber yields by decreasing cumulative light interception without affecting LUE. No differences in LUE between cultivars or cultivar classes were detected. Therefore, the maintenance of green leaf area is important when breeding potatoes for optimal performance in the presence of late blight. The results support the hypothesis that the correlation between lateness and reported resistance of potato cultivars is due to the vigorous foliage growth of late cultivars.     

The Effect of Nitrogen Rate on Vine Kill,Tuber Skinning Injury,Tuber Yield and Size Distribution,and Tuber Nutrients and Phytonutrients in Two Potato Cultivars Grown for Early Potato Production

Early potatoes are typically produced using less nitrogen than a full season potato crop as high rates of nitrogen may delay tuber set and lead to excessive vine growth that is difficult to terminate prior to harvest. Bintje and Ciklamen potato cultivars were grown with preplant soil nitrogen levels of 34 to 38, 67, and 101 kg N ha^-1 in 2013 and 2014 near Paterson, Washington. Nitrogen rate had little impact on the number of tubers and stems per plant of both cultivars, but increasing nitrogen rate tended to increase leaf area of both cultivars. Vine desiccation of Bintje with diquat was less complete as nitrogen rate increased, while Ciklamen vine kill was reduced by higher nitrogen in 1 of 2 years. Tuber skinning injury, tuber weight loss, and tuber size distribution were not affected by nitrogen rate. Tuber skinning injury and tuber weight loss were reduced in both cultivars by harvesting at 4 weeks after initial vine kill compared to harvesting at 2 weeks after vine kill. Total tuber yield was lower for both Bintje and Ciklamen in 1 of 2 years at the 34 to 38 kg N ha^-1 rate. Tuber nitrogen and zinc levels tended to increase with increasing nitrogen rates, while most other nutrients, vitamin C, total phenolics, and antioxidant capacity showed little response. It appears that 67 kg N ha^-1 provides adequate nitrogen to produce a good tuber set and yield of small tubers while not producing excessive vine growth that may be more difficult to kill.     

Effects of Toxic Levels of Aluminium on Seedling Parameters of Rice under Hydroponic Culture

The presence of Al in the rhizosphere of rice in acid soil restricts root growth and significantly reduces crop productivity.In this study,the effects of Al(30,60 and 90μg/mL)on seedling root growth,number of primary roots per seedling,seedling shoot length,number of leaves per seedling,seedling fresh weight,and seedling dry weight were studied.Rice genotypes were classified into three different classes,namely,tolerant,moderately tolerant,and susceptible,based on root tolerance index.The method of hydroponic culture was modified,and elaborated in the text.Toxic levels of Al in nutrient solution significantly decreased seedling root growth,number of primary roots,seedling shoot length,number of leaves per seedling,seedling fresh weight,and seedling dry weight.Few genotypes showed longer root length at 30μg/mL Al in nutrient solutions compared with the control.High levels of Al in nutrient solutions were highly toxic for rice seedlings.Based on root tolerance index,Radhunipagal,Gobindobhog,Badshabhog,Kalobhog,UBKVR-11,UBKVR-16,UBKVR-18,Khasha and IVT4007-B were classified as tolerant genotypes,and these genotypes may be used as donors for breeding of Altoxicity tolerance.     

Relationship between Deep Root Distribution and Root Penetration Capacity Estimated by Pot Experiments with a Paraffin and Vaseline Layer for Landraces and Recent Cultivars of Wheat

Root growth into deep soil is an important factor for stable production in wheat under drought conditions. Root penetrating capacity (RP) shown by pot experiments with a paraffin-Vaseline layer (PV layer) may be a useful indicator estimating deep rooting ability of wheat genotypes. Previously, we identified genotypes of durum wheat (Triticum turgidum L. var. durum) and bread wheat (T. aestivum L.) with diverse RP by the pot experiments. In this study, we investigated the root distribution of three Ethiopian landraces of durum wheat with high RP, three recent cultivars of durum wheat with low RP and one Japanese cultivar of bread wheat ‘Haruyutaka’ with low RP using: (1) pots with a PV layer, (2) root boxes, (3) artificial field and (4) a normal field to analyze the relationship between RP estimated by pot experiment and root development in the field. In the pot experiments, RP was evaluated by the number of roots penetrating through the PV layer (NRP). In the root-box and field experiments, the root distribution was evaluated by the number of roots on the vertical surface of soil as the root frequency (RF: root number cm-¹ soil surface). Ethiopian landraces had a significantly larger NRP than recent cultivars in the pot experiment. The root box and field experiments showed that Ethiopian landraces tended to have a higher RF than recent cultivars in deep soil layer. We concluded that RP estimated by pot experiments with a PV layer is a useful indicator of deep rooting ability under field conditions.     

Potato Crop Response to Genotype and Environment in a Subtropical Highland Agro-ecology

Potato response to environment, planting date and genotype was studied for different agro-ecological zones in Lesotho. Field experiments were conducted at four different sites with altitudes ranging from 1,655 to 2,250 m above sea level during the 2010/2011 and 2011/2012 summer growing seasons. Treatments consisted of three cultivars that varied in maturity type, two planting dates and four sites differing in altitude and weather patterns. Various plant parts were measured periodically. To understand and quantify the influence of abiotic factors that determine and limit yields, the LINTUL crop growth model was employed which simulated potential yields for the different agro-ecological zones using weather data collected per site during the study period. Observed actual crop yields were compared with model simulations to determine the yield gap. Model simulations helped to improve our understanding of yield limitations to further expand potato production in subtropical highlands, with emphasis on increasing production through increased yields rather than increased area. Substantial variation in yield between planting date, cultivar and site were observed. Average tuber dry matter (DM) yields for the highest yielding season were above 7.5 t DM ha^?1 or over 37.5 t ha^?1 fresh tuber yield. The lowest yield obtained was 2.39 t DM ha^?1 or 12 t ha^?1 fresh tuber yield for cultivar Vanderplank in the 2011/2012 growing season at the site with the lowest altitude. Modelled potential tuber yields were 9–14 t DM ha^?1 or 45–70 t ha^?1 fresh yield. Drought stress frequently resulted in lower radiation use efficiencies and to a lesser degree harvest indices, which reduced tuber yield. The site with the lowest altitude and highest temperatures had the lowest yields, while the site with the highest altitude had the highest yields. Later maturing cultivars yielded more than earlier maturing ones at all sites. It is concluded that the risk of low yields in rain-fed subtropical highlands can be minimised by planting late cultivars at the highest areas possible as early as the risks of late frosts permit.     

Two strategies for achieving higher yield under phosphorus deficiency in winter wheat grown in field conditions

Under field conditions, phosphorus (P) deficiency reduces wheat yield by affecting different yield components. However, the physiological strategies by which wheat genotypes with different yield structures respond to low-P stress are not clear. In the present study, we investigated tiller, floret, and root biomass, and P uptake and remobilization at two levels of P under field conditions in three winter wheat genotypes with different yield structures and P-efficiencies. Results showed that P-efficient cultivars CA9325 and ND139 got higher yield and total P accumulation than P-inefficient ND3291 at low-P, but not at normal P treatment. However, both the P-efficient wheat cultivars tend to have the same advantageous yield components at both high P and low-P stress. CA9325, a large-eared genotype, developed more fertile florets, and therefore had more grains at low-P stress. Increasing the number of grains formed a large sink for P demand during the grain-filling stage. Correspondingly, this genotype developed large roots for sustaining post-anthesis P uptake. ND139, a multi-eared genotype, developed many more tillers at low-P stress, and formed more ears at maturity. P from infertile tillers was probably reutilized by the surviving tillers to ensure floret development. Correspondingly, the contribution of pre-anthesis P uptake in ND139 and subsequent remobilization of P to the grains was higher. It was found that larger root rather than higher root activity was the determinant factor in efficient pre-anthesis P uptake in ND3291 and efficient post-anthesis P uptake in CA9325. It is concluded that increasing wheat yield at low soil P availability can be realized by either increasing ears per plant or increasing grains per ear through crop management or breeding.     

Root size,distribution and soil water depletion as affected by cultivars and environmental factors

Field experiments were carried out in two parts at the Luancheng Experimental Station in the North China Plain. Part I, which was a continuous experiment that ran from 1990 to 2008, investigated the change of grain yield, evapotranspiration (ET), root size and soil water utilization of winter wheat (Triticum aestivum L.) under three water regimes (rain-fed, two and four irrigation applications). Part II used 10 cultivars released from 1970 to 2000 that were grown under the same condition for two seasons (2005/2006 and 2006/2007) to compare root size, root:shoot ratio and soil water depletion (SWD). The results of testing in Part I showed that the yield and ET of winter wheat gradually increased from 1990 to the present. There was no consistent change in total root length (TRL) over time. The difference in root size among seasons and irrigation treatments mainly occurred in the upper soil profile, where the root length density (RLD) was greater. No direct relationship was found between root size and soil water use. Thus, TRL was not a factor that indicated the water extracting capacity of crops. The results from Part II revealed that the seasonal ET of earlier released cultivars (ERC) was similar to that of recently released cultivars (RRC) under well-watered conditions. However, ET was slightly increased from ERC to RRC under water deficit conditions, indicating high soil water depletion by RRC. TRL decreased from ERC to RRC and was significantly correlated with plant height. The breeding of winter wheat that reduced plant height not only increased harvest index, but also reduced root size, resulting in a smaller root:shoot ratio. The reduction in TRL from ERC to RRC mainly occurred in the top soil profile. The results from both tests indicate that TRL is not a factor that determines soil water use; rather, the distribution of RLD along the soil profile plays more important role in soil water utilization. Smaller root size in the upper soil layer did not affect soil water uptake, and might be more economical in terms of production efficiency.     

Root System Development Including Root Branching in Cuttings of Cassava with Reference to Shoot Growth and Tuber Bulking

Summary

In spite of the important role it plays for water and nutrient acquisition, information on the root system development in cassava (Manihot esculenta Grantz) is limited. To examine the root length and branching pattern with reference to shoot growth and tuber bulking, we grew cassava plants in containers under natural climatic conditions in the southern end of Sumatra Island, Indonesia. One 20-cm length cutting of cassava (cv. Ardira IV) was planted in either a plastic bucket or a wooden box. The containers, which were filled with heavy clay soil, had different sizes depending on the growing period. At 30, 60, 100, 140, 180, and 270 days after planting (DAP), both the shoot and roots were sampled for quantitative analysis. The dry weight of both shoot and roots increased rapidly with the leaf area. The axile root number, however, decreased from 60 to 140 DAP as a result of the abscission of roots emerging from the basal part of the cutting during tuber bulking. The total root length reached its maximum at 60 DAP and significantly decreased thereafter because of decay and decomposition during tuber bulking. On the other hand, the root branching either increased the branching order or retained it, as determined from a topological point of view. The root branching during the later growing period compensated for the decrease in total root length and contributed to maintain a sufficient root surface area. The surviving roots with a well-developed branching pattern could absorb water and nutrients essential for tuber bulking.