Partial Root Drying Effects on Biomass Production in Brassica napus and the Significance of Root Responses

Partial root drying (PRD) has been shown to stimulate stomatal-closure response and improve water-use efficiency and thus biomass production and grain yield under water deficiency. While most studies focus on above-ground responses to PRD, we examined how root responses contributed to effects of partial root drying. In particular, in two experiments with oilseed rape (Brassica napus L.) we investigated whether roots were able to forage for patchily distributed water, and how this affected plant growth compared with uniform watering and alternate watering (in which different parts of the roots receive water alternately). The first pot experiment was carried out in the greenhouse and the second outside under a rain-shelter in which also the watering amount was varied. The results indicate that B. napus roots were able to forage for fixed water patches by selective root placement. In the first experiment with small plants, root foraging was equally effective as enhanced water-use efficiency under alternate watering. Both treatments resulted in about 10% higher shoot biomass compared with uniform watering. Alternate watering generally outperformed uniform watering in the second experiment, but the success depended on the time of harvest and the water supply level. Measurements indicated that only the alternate watering regime effectively reduced stomatal conductance, but lead to a higher shoot biomass only under more severe (50%) rather than under milder water deficiency (70% of a well watered control). Water deficiency strongly reduced leaf initiation rates and leaf sizes in B. napus, but for a given level of water supply the supply pattern (uniform control, fixed patchy or alternate watering) hardly influenced these growth parameters. Although also in the second experiment, the plants selectively placed their roots in the wet parts of the pot, root foraging was not as effective as in the first experiment. Possible reasons for these discrepancies are discussed as well as their implications for the application of PRD effects for crop growth.     

Root plasticity maintains growth of temperate grassland species under pulsed water supply

Background and aims

The frequency of rain is predicted to change in high latitude areas with more precipitation in heavy, intense events interspersed by longer dry periods. These changes will modify soil drying cycles with unknown consequences for plant performance of temperate species.

Methods

We studied plant growth and root traits of juveniles of four grasses and four dicots growing in a greenhouse, when supplying the same total amount of water given either regular every other day or pulsed once a week.

Results

Pulsed water supply replenished soil moisture immediately after watering, but caused substantial drought stress at the end of the watering cycle, whereas regular watering caused more moderate but consistent drought. Grasses had lower water use efficiency in the pulsed watering compared to regular watering, whereas dicots showed no difference. Both grasses and dicots developed thinner roots, thus higher specific root length, and greater root length in the pulsed watering. Growth of dicots was slightly increased under pulsed watering.

Conclusions

Temperate species coped with pulsed water supply by eliciting two responses: i) persistent shoot growth, most likely by maximizing growth at peaks of soil moisture, thus compensating for slower growth during drought periods; ii) plasticity of root traits related to increased resource uptake. Both responses likely account for subtle improvement of growth under changed water supply conditions.     

Grain Yield of Pennisetum americanum Adjusts to Nitrogen Supply by Changing Rates of Grain Filling and Root Uptake of Nitrogen

Coaldrake, P. D., Pearson, C. J. and Saffigna, P. G. 1987. Grainyield of Pennisetum americanum adjusts to nitrogen supply bychanging rates of grain filling and root uptake of nitrogen.–J.exp. Bot 38: 558–566. Pearl millet (Pennisetum americanum(L.)Leeke) was grown in containers at three constant rates of nitrogensupply or with the nitrogen supply increased from the lowestto the highest rate during panicle differentiation or at anthesis.We measured the rate and duration of nitrogen and dry weightgain by individual grains and nitrogen (¹⁵N) uptake by rootsand its distribution during grain filling. The total amountsof nitrogen and dry weight in all grain per plant at the lowestnitrogen supply were 8% and 14% respectively of plants growncontinuously at the highest rate of nitrogen. This was becauselow rates of nitrogen supply reduced grain number, mean grainweight and the nitrogen content of each individual grain. Theamino acid composition of the grain protein was affected onlyslightly by nitrogen treatments. Rates of grain growth were sensitive to nitrogen supply whereasthe duration of nitrogen movement to the grain was not. Nitrogenuptake by roots continued throughout grain filling; rates ofuptake per g root in plants given least nitrogen were one-halfthose of plants given the highest amount of nitrogen. A changefrom lowest to highest nitrogen supply at panicle differentiationincreased the uptake of nitrogen by roots and the rates of growthof individual grains, to the rates observed in plants whichhad been supplied continuously with the highest nitrogen. Whenthe change in supply was made at anthesis there was rapid movementof nitrogen into the plant but this was not translated intomore rapid grain growth. Key words: Nitrogen supply, Pennisetum americanum, grain yield, root uptake     

Root Growth Inhibitors from Root Cap and Root Meristem of Zea mays L.

A micro-assay based on the growth inhibition of root segmentsof the seminal roots of Zea mays has been used to investigatethe root-growth-inhibiting substances in root caps and meristemsrespectively of the roots of Zea mays. This micro-assay is sensitiveto 50 pg of IAA or less. Paper chromatography of the acid fractionof methanolic extracts shows the presence of one main inhibitorin root caps and a different main inhibitor in root meristems.Neither is IAA, whose presence in meristems is sometimes indicatedby small inhibitions (or stimulations) at the characteristicRf of IAA. A Commelina leaf-epidermis assay shows the presenceof one stomata-closing ABA-like substance in root caps and onein meristems, one corresponding in Rf to the main root-growthinhibitor from the root cap. The implications of these findingsfor the geotropic responses of roots is briefly discussed.     

Changes in the Root System of Wheat Seedlings Following Root Anaerobiosis I. Anatomy and Respiration in Triticum aestivum L.

Anatomical changes in roots of wheat seedlings (Triticum aestivumL. cv. Hatri) following oxygen deficiency in the rooting mediumwere investigated. The response of the plant to stress was testedat a very early developmental stage when the first adventitiousroots had just emerged. In order to analyze the adaptation ofdifferent roots, respiration rates of the roots 1–3 and4–n were compared with the respiration rates of the totalroot system. Oxygen deficiency was induced either by flushingnutrient solution with nitrogen or flooding of sand. In contrast to plants grown in well aerated media, both stressvariants led to a significant increase of the intercellularspace of the root cortex in seminal and first adventitious roots.Radial cell enlargement of cortical cells near the root tip,cell wall thickenings in flooded sand cultures and an increasein phloroglucinol-stainable substances were found to be furtherindicators of low oxygen supply. The roots 4–n which were promoted in growth under hypoxiashowed higher respiration rates; hence the total root respirationwas not restricted. Triticum aestivum L. cv. Hatri, wheat, roots, anatomy, anaerobiosis, stress, root respiration, intercellular space     

The Rate of Growth and Partitioning of Assimilates in Young Grass Plants: A Mathematical Model

A model describing the increase in weight with time of younggrass plants is formulated. The parameters are the relativegrowth rates of the root and shoot systems; k, the ratio ofthe relative growth rate of the root system relative to thatof the shoot system; b, the weight of the root system when thatof the shoot system is unity, and u the rate of increase inweight of the whole plant per unit of shoot system per unitof time, k and b are the constants in the allometric formula,r = bs^k where r and are the weights of the root and shoot systems.The model enables the effect of changes in the distributionof assimilates between the root and shoot systems upon the rateof growth of the plant to be assessed. Data from a number ofexperiments are analysed in this manner and the significanceof the results discussed.     

Further Observations on Root Contraction

Further observations, by light and electron microscopy, on theanatomy of root contraction in Hyacinthus orientalis cvs. arereported, together with some comparisons with Narcissus andGladiolus. It is maintained that root contraction in these andsome other species is essentially a growth process, as previousinvestigators have shown, and that an understanding of the re-directedgrowth of the cortical cells which brings it about may be basedon the specialized structural features of their cell walls,here described. Physical, anatomical and comparative reasonsare given for our belief that the alternative explanation ofthe contractile process in Gladiolus which relates it to cellcollapse resulting from transpirational ‘pull’ isuntenable. anatomy, contraction, Hyacinthus, Narcissus, Gladiolus, root     

Adaptation and Functional Balance Between Shoot and Root Activity of Sunflower Plants Grown at Different Root Temperatures

Helianthus annuus L. plants were grown with the shoots at normalair temperature and with the roots in nutrient solution at 10,20 or 30 °C. The higher the root temperature the greaterthe growth of the leaves, resulting in higher production ofphotosynthates. Irrespective of growth conditions an equilibriumwas established between the maintenance respiratory activityof shoot and roots. A constant proportion of the photosynthateproduced was used in respiration. The results are discussedin relation to a thermodynamic theory of stability of biologicalsystems. It is suggested that changes in energy partition betweenmaintenance and growth, and then in relative growth rates ofshoots and roots during an adaptation period, represent a majorhomeostatic mechanism. shoots, roots, maintenance respiration, growth, relative growth rate, respiration, adaptation, sunflower, Helianthus annuus L.     

Root Distribution, Growth, Respiration, and Hydraulic Conductivity for Two Highly Productive Agaves

Cultivated Agave mapisaga and A. salmiana can have an extremelyhigh above-ground dry-weight productivity of 40 Mg ha^–1yr^–1. To help understand the below-ground capabilitiesthat support the high above-ground productivity of these Crassulaceanacid metabolism plants, roots were studied in the laboratoryand in plantations near Mexico City. For approximately 15-year-oldplants, the lateral spread of roots from the plant base averaged1.3 m and the maximal root depth was 0.8 m, both considerablygreater than for desert succulents of the same age. Root andshoot growth occurred all year, although the increase in shootgrowth at the beginning of the wet season preceded the increasein growth of main roots. New lateral roots branching from themain roots were more common at the beginning of the wet season,which favoured water uptake with a minimal biomass investment,whereas growth of new main roots occurred later in the growingseason. The root: shoot dry weight ratio was extremely low,less than 0.07 for 6-year-old plants of both species, and decreasedwith plant age. The elongation rates of main roots and lateralroots were 10 to 17 mm d^–1, higher than for various desertsucculents but similar to elongation rates for roots of highlyproductive C₃ and C₄ agronomic species. The respiration rateof attached main roots was 32 µmol CO₂ evolved kg^–1dry weight s^–1 at 4 weeks of age, that of lateral rootswas about 70% higher, and both rates decreased with root age.Such respiration rates are 4- to 5-fold higher than for Agavedeserti, but similar to rates for C₃ and C₄ agronomic species.The root hydraulic conductivity had a maximal value of 3 x 10^–7ms^–1 MPa^–1 at 4 weeks of age, similar to A. deserti.The radial hydraulic conductivity from the root surface to thexylem decreased and the axial conductivity along the xylem increasedwith root age, again similar to A. deserti. Thus, although rootsof A. mapisaga and A. salmiana had hydraulic properties perunit length similar to those of a desert agave, their highergrowth rates, their higher respiration rates, and the greatersoil volume explored by their roots than for various desertsucculents apparently helped support their high above-groundbiomass productivity Key words: Crassulacean acid metabolism, productivity, root elongation rate, root system, water uptake     

The Effects of Infection by Rust (Puccinia lagenophorae Cooke) on the Growth of Groundsel (Senecio vulgaris L.) Cultivated under a Range of Nutrient Concentrations

Groundsel (Senecio vulgaris L.), healthy or infected with therust fungus Puccinia lagenophorae Cooke, was grown at a rangeof nutrient concentrations in sand culture. There were statisticallysignificant interactions between the effects of infection andnutrient supply upon the dry weights of stems, leaves, rootsand reproductive tissues, leaf area and cumulative capitulumproduction. This interaction occurred since infection causedsignificant inhibitions of growth only at moderate or high nutrientconcentrations. At low concentrations rusted plants were similarto or slightly larger than controls. Both in controls and rustedplants root: shoot ratios increased as nutrient supply declined.The ratio of root: shoot dry weight was consistently reducedby infection whilst root length: leaf area ratio was relativelyunchanged. More detailed investigations confirmed that infection had littleeffect on plant growth under nutrient deficient conditions despitesuppression of the host's ability to increase root: shoot ratiosin response to nutrient stress. This reflected the inhibitionof relative growth rates in rusted plants at high but not lownutrient concentrations, which in turn reflected reduced netassimilation rates (NAR). Increases in leaf-area ratio (LAR)often ameliorated the decline in NAR in rusted plants. Senecio vulgaris L., Puccinia lagenophorae Cooke, nutrient deficiency, growth, root: shoot ratio     

A Study of the Nitrogen and Water Factors in Competition with Agropyron repens (L.) Beauv.

From a consideration of the possible relationships between thelevel of a nutrient in the soil, the depletion due to a competitor,and the response of a test plant, it is argued that a validcriterion for the implication of a nutrient factor in root competitionis that some growth response of the test plant to competitionshould be less at higher levels of supply of that nutrient thanat lower levels. Young test plants of Impatiens parviflora were transplantedinto pans of soil in which stands of Agropyron repens were growing.Two levels of nitgrogen and gtwo rates of watering were appliedin a multifactorial design, and from half of the test plantsroot competition was excluded, so that their responses to rootcompetition could be measured under the various nitrogen andwatering treatments. A preliminary experiment hadf shown, and the competition experimentconfirmed, that the optimum nitrogen level for root growth andNet Assimilation Rate was less than the higher level employed,whereas this level was not sufficient for maximum leaf-growth.In agreement with this, the depressions of relative root growth-rateand Net Assimilation Rate caused by competition at the lowernitrogen level were significantly reduced at the higher level,but for relative leaf-growth this reduction did not reach significance.The effects of competition were reduced by the higher rate ofwstering, and for relative leaf growth-rate this interactionwas highly significant. It is deduced that competition from A. repensfinvolves bothnitrogen and water, and that water is probably the more importantfactor under normal conditions. There is no need to postulkateany toxic root product to account for the experimentalo results.Compared with competition above ground, it appeared that rootcompetition could have an effect on the relative growth-rateof I. parviflora about as great as a reduction of light intensityfrom full to 5–10 per cent. of summer daylight.     

Distinct Influence of Root and Shoot Temperature on Nitrogen Fixation by White Clover

White clover (Trifolium repens L.) was grown in controlled environmentsto determine the distinct effects of root and shoot temperatureon the accumulation of total and fixed (¹⁵ N dilution) nitrogenat two levels of nitrate (10 and 75 mM). Nitrogen fixation(BNF) showed a positive response to higher shoot temperature(23 vs. 13 C day temperature), irrespective of whether or notroot temperature was increased in parallel. Low root temperature(5 C) caused a marked reduction in the accumulation of totalnitrogen at both nitrate levels, and led to a lower proportionof N derived from BNF. The temperature response of BNF was attributedfor the major part to an adaptation to the demand for fixedN. It is therefore concluded that BNF is not primarily responsiblefor the reduced clover growth at low temperatures. White clover, Trifolium repens L., temperature, nitrogen fixation, nitrate, root, shoot     

Root Morphology and Nitrogen Uptake of Maize Simultaneously Supplied with Ammonium and Nitrate in a Split-root System

We studied the response of maize (Zea mays L. cv. Anjou 256)to a simultaneous, but separated supply of ammonium and nitrate(localized supply, LS). A split-root system was used to supplyhalf of the roots with ammonium and the other half with nitrate.A homogeneously distributed supply of both nitrogen forms (HS)was the control treatment. Seedlings were grown for 12 d fromthe two-leaf to the three-leaf stage in hydroponics at threepH levels (4, 5·5 and 7). The total N concentration was3 mol m^-3. The split-root system was established by removingthe seminal root system and using only four nodal roots perplant. Total root length and root surface area were recordedautomatically with a modified Delta- T area meter. Other morphologicalroot traits (such as main axis length and diameter, number,density, and length of laterals) were recorded manually. Uptakeof ammonium and nitrate was measured by the depletion of thenutrient solution. As compared with LS, HS was superior in shootand root DM, total root length and root surface area, ammoniumand nitrate uptake and shoot nitrogen concentration, irrespectiveof pH level. This indicates that, also under field conditions,mixed ammonium and nitrate fertilization is only beneficialto plant growth if both N forms are evenly distributed in thesoil. At both HS and LS, ascending pH increased the ammonium:nitrateuptake ratio. At LS, declining pH induced a considerable shiftin the distribution of root DM, root length, and root surfacearea the nitrate-fed compartment.Copyright 1993, 1999 AcademicPress Maize, Zea may L., ammonium, nitrate, pH, root morphology, split-root     

Cell Cycle Duration in the Root Meristem of Sonoran Desert Cactaceae as Estimated by Cell-flow and Rate-of-cell-production Methods

Slow rates of cactus growth in the Sonoran Desert and high productivityof some Cactaceae under cultivation suggest that relativelylow growth rates are not the consequence of a long cell divisioncycle but of short optimal periods for growth and adverse environmentalfactors. To verify this hypothesis, the duration of the celldivision cycle (T)in the root apical meristem of seedlings ofthree sympatric species from the Sonoran Desert [Ferocactuspeninsulae(F. A. C. Weber) Britton & Rose ‘Townsendianus’(Britton & Rose) N. P. Taylor, stat. nov.,Stenocereus gummosus(Engelm.)Gibson & Horak andPachycereus pringlei(S. Watson) Britton& Rose] was estimated with the rate-of-cell-production (RCP)and the cell-flow (colchicine) methods. Both methods were appliedduring the steady-state growth phase, which was relatively shortin the first two species because of the determinate patternof root growth. The RCP method permitted estimation ofTin eachroot individually. Durations of the cell division cycle wereinversely proportional to the rate of root growth (r²rangedfrom 0.42 to 0.88,P<0.05).T,determined by the cell-flow method,ranged from 14.4 to 19.3 h in these species and was within thesame range asTdetermined by the RCP method. The averageTdeterminedby the RCP method was 67 to 75% of that determined by the cell-flowmethod. Results obtained with both methods are compared andanalysed. The proposed hypothesis appears to be correct, indicatingthat these species can be more productive under cultivationthan in the wild due to the relatively short duration of thecell division cycle. Adaptive features of these findings arealso considered.Copyright 1998 Annals of Botany Company Cactaceae, cell division cycle, root growth, root meristem, Sonoran Desert     

Vegetable Plant Part Relationships. II. A Quantitative Hypothesis for Shoot/Storage Root Development

A simple quantitative formulation of the concept of the controlof partitioning of assimilated carbon by the behaviour of plantcomponents as competing sinks is developed. An equation, In s = + In r—t, relating shoot (s) and storage root (r) d. wts, and the lengthof growth period (t), is constructed by considering possiblefates of imported assimilates into different plant parts. Thevalues of the equations' parameters depend on the relative sinkactivities of the plant parts, tissue respiration rates andinitial weights of plant components. The equation closely fitteddata collected from a number of carrot and beet experimentsin which planting density had been varied. Estimates of shootand storage root maintenance respiration rates, derived fromthe parameter , were of the correct order of magnitude. Othersets of experimental data are also discussed in the light ofpredictions of the theory and possible uses and extensions ofthis approach to assimilate partitioning are briefly discussed. Daucus carota L., Beta vulgaris, carrot, red beet, partition of assimilated carbon, maintenance respiration, storage root     

Adventitious Root Formation in Veronica Spp.

Species of the genus Veronica differ in habitat preferences,growth form and in adventitious root production. The annualspecies rarely or never produce adventitious roots in intactplants in the field but some, for example V. persica and V.arvensis will root vigorously from single node stem segmentsin culture. Others, such as V. agrestis require the presenceof IAA for substantial levels of root formation to occur incultured stem segments. Veronica hederifolia cuttings rarelyproduce roots. Stem cuttings of the perennial species, in general,rooted more vigorously than those of annual plants. Both V.fihiformis and V. serpyllifolia root very strongly. The position of root production from the stem cuttings differedfrom species to species. Roots arose either from the node, theregion of the base or at some intermediate point. Veronica arvensis,V. chamaedrys and V. persica rooted mainly from the basal regionwhereas V. filiformis rooted mainly from the node. Veronicaserpyllifolia cuttings rooted at both of these locations. Veronica filiformis, a perennial species that is infertile inBritain, produces root primordia in intact plants at nodes whichare close to the shoot apex. Thus, even very young stem segmentshave ‘preformed’ root primordia. For this reason,detached stem segments of V. filiformis root very rapidly andthis probably has been of great significance in its successfulinvasion and spread in lawns and short turf areas. Veronica spp., adventitious roots, indol-3-ylacetic acid, root primordia, vegetative reproduction     

Episodic Growth and Relative Shoot: Root Balance in Loblolly Pine Seedlings

Leaf, root and stem systems of loblolly pine seedlings are characterizedby a seasonal periodicity in growth, during which they alternatein spurts of activity. Despite this periodicity, the allometriccoefficient describing the ratio of the relative growth ratesof leaf to root remains constant for at least the first twoyears of development. In part, constancy results from the inabilityof variation in the allocation of growth increment during briefperiods to change a pre-existing structure accumulated overthe life of the seedling. In addition, alternating periods ofleaf, root and stem growth may represent the action of feedbackmechanisms which operate to maintain an adaptive balance betweenorgan systems. Pinus taeda L., loblolly pine, allometric coefficient, homeostatic control of growth