Patterns of variability in the diameter of lateral roots in the banana root system

The relative importance of root system structure, plant carbon status and soil environment in the determination of lateral root diameter remains unclear, and was investigated in this study. Banana (Musa acuminata) plants were grown at various moderate levels of soil compaction in two distinct experiments, in a field experiment (FE) and in a glasshouse experiment (GE). Radiant flux density was 5 times lower in GE. The distribution of root diameter was measured for several root branching orders. Root diameters ranged between 0.09 and 0.52 mm for secondary roots and between 0.06 and 0.27 mm for tertiary roots. A relationship was found between the diameter of the parent bearing root and the median diameter of its laterals, which appears to be valid for a wide range of species. Mean lateral root diameter increased with distance to the base of the root and decreased with branching density [number of lateral roots per unit length of bearing root (cm(-1))]. Typical symptoms of low light availability were observed in GE. In this case, lateral root diameter variability was reduced. Although primary root growth was affected by soil compaction, no effects on lateral root diameter were observed.     

Relationships between root diameter,root length and root branching along lateral roots in adult,field-grown maize

Background and Aims Root diameter, especially apical diameter, plays an important role in root development and function. The variation in diameter between roots, and along roots, affects root structure and thus the root system’s overall foraging performance. However, the effect of diameter variation on root elongation, branching and topological connections has not been examined systematically in a population of high-order roots, nor along the roots, especially for mature plants grown in the field.Methods A method combining both excavation and analysis was applied to extract and quantify root architectural traits of adult, field-grown maize plants. The relationships between root diameter and other root architectural characteristics are analysed for two maize cultivars.Key Results The basal diameter of the lateral roots (orders 1–3) was highly variable. Basal diameter was partly determined by the diameter of the bearing segment. Basal diameter defined a potential root length, but the lengths of most roots fell far short of this. This was explained partly by differences in the pattern of diameter change along roots. Diameter tended to decrease along most roots, with the steepness of the gradient of decrease depending on basal diameter. The longest roots were those that maintained (or sometimes increased) their diameters during elongation. The branching density (cm^–1) of laterals was also determined by the diameter of the bearing segment. However, the location of this bearing segment along the mother root was also involved – intermediate positions were associated with higher densities of laterals.Conclusions The method used here allows us to obtain very detailed records of the geometry and topology of a complex root system. Basal diameter and the pattern of diameter change along a root were associated with its final length. These relationships are especially useful in simulations of root elongation and branching in source–sink models.     

The effect of Tiron, a water soluble radical scavenger, on growth, morphology and alkaloid content of adventitious roots in Atropa belladonna

 The effect of Tiron (disodium 1,2-dihydroxybenzene-3,5-disulfonate) on the growth, morphology and alkaloid content of adventitious roots in Atropa belladonna was investigated. High concentrations of Tiron had an inhibitory effect on growth of the root. The appearance of cultured roots was significantly changed from rough roots accompanied with callus-like tissue in control cultures to fine roots without callus formation. Alkaloid content was drastically increased by the addition of 1 mM Tiron to the medium. The influence of NAA, which has an inhibitory effect on alkaloid production, was partially restored by Tiron treatment, indicating that this radical scavenger may affect the production of alkaloids through modulation of the mode of action of auxin. Glutathione content of the root was not influenced by Tiron. Received: 3 June 1999 / Revision received: 28 September 1999 / Accepted: 30 September 1999     

Domestication and crop physiology: roots of green-revolution wheat

BACKGROUND AND AIMS: Most plant scientists, in contrast to animal scientists, study only half the organism, namely above-ground stems, leaves, flowers and fruits, and neglect below-ground roots. Yet all acknowledge roots are important for anchorage, water and nutrient uptake, and presumably components of yield. This paper investigates the relationship between domestication, and the root systems of landraces, and the parents of early, mid- and late green-revolution bread wheat cultivars. It compares the root system of bread wheat and 'Veery'-type wheat containing the 1RS translocation from rye. METHODS: Wheat germplasm was grown in large pots in sand culture in replicated experiments. This allowed roots to be washed free to study root characters. KEY RESULTS: The three bread wheat parents of early green-revolution wheats have root biomass less than two-thirds the mean of some landrace wheats. Crossing early green-revolution wheat to an F(2) of 'Norin 10' and 'Brevor', further reduced root biomass in mid-generation semi-dwarf and dwarf wheats. Later-generation semi-dwarf wheats show genetic variation for root biomass, but some exhibit further reduction in root size. This is so for some California and UK wheats. The wheat-rye translocation in 'Kavkaz' for the short arm of chromosome 1 (1RS) increased root biomass and branching in cultivars that contained it. CONCLUSIONS: Root size of modern cultivars is small compared with that of landraces. Their root system may be too small for optimum uptake of water and nutrients and maximum grain yield. Optimum root size for grain yield has not been investigated in wheat or most crop plants. Use of 1RS and similar alien translocations may increase root biomass and grain yield significantly in irrigated and rain-fed conditions. Root characters may be integrated into components of yield analysis in wheat. Plant breeders may need to select directly for root characters.     

Maximum axial growth pressures of the lateral roots of pea and eucalypt

Although lateral roots may contribute significantly towards growth and nourishment of plants, the mechanics of their elongation behaviour in strong soils is not well known. The aim of this study is to report maximum axial growth pressures (p) and maximum elongation rates (E) of the lateral roots of an annual herbaceous plant (pea) and a woody perennial (eucalypt). As such measurements have not been reported previously, measurements of P and E for lateral roots were compared with the primary roots of pea for which reports are widespread. Values of P were estimated from the measured maximum values of axial force and root diameter on single, intact roots of seedlings in the laboratory. Additional measurements of both P and E were made for the lateral roots of pea when the growth of the remaining root axes was stopped (with removal of tips) to determine the overall effects of root-growth-inhibition on P and E of single roots.Values of P and E for lateral roots of pea were significantly greater than those for the lateral roots of eucalypt. Although root diameter for the primary roots of pea were similar to those for the lateral roots of eucalypt, the former exerted nearly twice as much pressure as the latter. The lateral roots of pea elongated significantly slower than the primary roots; however, P of lateral roots was significantly lower than the primary roots when elongation of all other roots was inhibited during the measurements. Production and/or development of lateral roots increased when elongation of the remaining roots (both primary and lateral roots) of pea seedlings was restricted due to the removal of root tips and exposure of one of the lateral roots to high strength. In general, maximum axial force exerted by primary and lateral roots was similar for roots of <1 mm diameter. However, primary roots exerted greater maximum axial force than the lateral roots when root diameter was >1 mm. As axial pressure of lateral roots was independent of root diameter, thickening of root tips is less likely to assist penetration of lateral roots in strong soils.     

玉米初生根和不同层次次生根的直径和大量元素含量的比较分析

对大田条件下玉米品种'掖单13'(Zea mays 'Yedan 13')和'丹玉13'(Z. mays 'Danyu 13')初生根(包括初生胚根和次生胚根)和不同层次次生根的直径及其中的N、P、K、Ca和Mg含量进行了比较分析.分析结果表明,'掖单13'和'丹玉13'初生根直径总体上小于次生根;从第1层至第7层次生根,基本上为发生时间越晚,根直径越大;而2个品种第7层至第9层次生根基部直径的变化趋势有一定的差异.'掖单13'和'丹玉13'初生根和不同层次次生根中N、P、K、Ca和Mg含量差异显著(P<0.05);'掖单13'初生根中的N、P、K、Ca和Mg含量分别为27.36、 3.75、 14.34、 10.19和5.94 g·kg~(-1), 不同层次次生根中的N、 P、 K、 Ca和Mg含量分别为4.31～8.99、 0.91～1.29、4.07～15.27、2.35～5.42和1.52～4.06 g·kg~(-1);'丹玉13' 初生根中的N、P、K、Ca和Mg含量分别为28.65、4.65、14.54、10.33和6.04 g·kg~(-1),不同层次次生根中的N、P、K、Ca和Mg含量分别为7.41～12.70、0.84～1.27、1.81～24.69、2.46～3.73和1.12～2.73 g·kg~(-1);2个品种初生根中N、P、Ca和Mg含量均最高,K含量也显著高于第1层至第7层次生根;总体上看,不同层次次生根中的N、Ca和Mg含量随发生时间的先后呈逐渐降低的趋势,P和K含量随发生时间的先后呈逐渐增加的趋势,但2个品种间有一定的差异.研究结果显示,次生根根直径的增大与其对地上部分的固持能力有关.     

Growth direction of nodal roots in rice: its variation and contribution to root system formation

The root system of a rice plant (Oryza sativa L.) consists of numerous nodal roots and their laterals. The growth direction of these nodal roots affects the spatial distribution of the root system in soil, which seems to relate to yield and lodging resistance. The growth angle of a nodal root varies with the type and timing of emergence of the nodal root. The body of a rice plant can be recognized as an integrated set of shoot units, each unit consisting of an internode with a leaf and several roots. Nodal roots formed at the apical part of a shoot unit often elongate horizontally, whereas those formed at the basal part of the shoot unit show various growth directions depending on both the growth stages of the plant and the environmental conditions. Moreover, nodal roots that emerge from the most basal shoot unit of a tiller are usually thick and grow downwards. External factors such as planting density and nitrogen application affect the growth direction of nodal roots, probably partly because of the changing tillering pattern of the shoot. In addition to the growth angle of nodal roots, size of nodal roots may be another important factor determining the spatial distribution of the root system in soil.     

Effects of 60-Hz electric fields on cellular elongation and radial expansion growth in cucurbit roots

Serial longitudinal and transverse sections were prepared from roots of Cucumis sativus and Cucurbita maxima that had been exposed/sham-exposed to 60-Hz electric fields for 0-2 days. Field exposures were selected to produce a 10-20% or a 70-80% growth inhibition in whole roots of both species. Cortical cell length and diameter were measured using a microscope and eyepiece micrometer; measurements were conducted "blind." In both species, inhibition of cellular elongation was associated with exposure to electric fields (EF). Cellular radial expansion was apparently unaffected by exposure to electric fields. The diameters of radially unexpanded or fully expanded C. sativus cortical cells were about 25-30% smaller than those of comparable cells in C. maxima roots. Previous studies of the relationship between rates of root growth and applied EF strength showed that the response thresholds of C. sativus and C. maxima differed by a similar relative amount. These results are consistent with the postulate that EF-induced effects in roots are elicited by induced transmembrane potentials.     

Microgeographic genome size differentiation of the carob tree, Ceratonia siliqua, at 'Evolution Canyon', Israel

BACKGROUND AND AIMS: We tested whether the local differences in genome size recorded earlier in the wild barley, Hordeum spontaneum, at 'Evolution Canyon', Mount Carmel, Israel, can also be found in other organisms. As a model species for our test we chose the evergreen carob tree, Ceratonia siliqua. METHODS: Genome size was measured by means of DAPI flow cytometry. KEY RESULTS: In adults, significantly more DNA was recorded in trees growing on the more illuminated, warmer, drier, microclimatically more fluctuating 'African' south-facing slope than in trees on the opposite, less illuminated, cooler and more humid, 'European' north-facing slope in spite of an interslope distance of only 100 m at the canyon bottom and 400 m at the top. The amount of DNA was significantly negatively correlated with leaf length and tree circumference. In seedlings, interslope differences in the amount of genome DNA were not found. In addition, the first cases of triploidy and tetraploidy were found in C. siliqua. CONCLUSIONS: The data on C. siliqua at 'Evolution Canyon' showed that local variability in the C-value exists in this species and that ecological stress might be a strong evolutionary driving force in shaping the amount of DNA.     

A simple model for nitrogen-limited plant growth and nitrogen allocation

BACKGROUND: and Aims In many studies of nitrogen-limited plant growth a linear relationship has been found between relative growth rate and plant nitrogen concentration, showing a negative intercept at a plant nitrogen concentration of zero. This relationship forms the basis of the nitrogen productivity theory. On the basis of empirical findings, several authors have suggested that there is also a distinctive relationship between allocation and plant nitrogen concentration. The primary aim of this paper is to develop a simple plant growth model that quantifies this relationship in mathematical terms. The model was focused on nitrogen allocation to avoid the complexity of differences in nitrogen concentrations in the different plant compartments. The secondary aim is to use the model for examining the processes that underlie the empirically based nitrogen productivity theory. METHODS: In the construction of the model we focused on the formation and degradation of biologically active nitrogen in enzymes involved in the photosynthetic process (photosynthetic nitrogen). It was assumed that, in nitrogen-limiting conditions, the formation of photosynthetic nitrogen is proportional to nitrogen uptake. Furthermore it was assumed that the degradation of photosynthetic nitrogen is governed by first-order kinetics. Model predictions of nitrogen allocation were compared with data from literature describing four studies of growth. Model predictions of whole plant growth were compared with the above-mentioned nitrogen productivity theory. KEY RESULTS: Allocation predictions agreed well with the investigated empirical data. The ratio of leaf nitrogen and plant nitrogen declines linearly with the inverse of plant nitrogen concentration. Nitrogen productivity is proportional to this ratio. Predictions for whole-plant growth were in accordance with the nitrogen productivity theory. CONCLUSIONS: The agreement between model predictions and empirical findings suggests that the derived equation for nitrogen allocation and its relationship to plant nitrogen concentration might be generally applicable. The negative intercept in the linear relationship between relative growth rate and plant nitrogen concentration is interpreted as being equal to the degradation constant of photosynthetic nitrogen.     

The relationships between static and dynamic variables in the description of root growth. Consequences for field interpretation of rooting variability

Field root investigations are often limited by the static nature of classical observations, resulting in the need to develop alternative methodologies that allow dynamic interpretation of root architecture variability on the basis of static measurements. The objectives of this work were (i) to evaluate the use of selected morphological indicators, namely root apical diameter (Da) and the length of the apical unbranched zone (LAUZ), in predicting primary and lateral root growth patterns in banana trees, (ii) to propose a field methodology for the assessment of root dynamics based on static measurements. Banana trees (Musa acuminata cv `Grande Naine') were grown in 5 rhizotrons as well as in field conditions, respectively on pouzzolane and Mollic Andosols. In rhizotrons, root growth analysis was carried out by reporting root elongation, Da and LAUZ, three times a week. In field conditions, 4 series of excavations were made at three-week intervals. Apart from root growth rate, measurements were the same as those in the rhizotrons. LAUZ was confirmed as a stable and good predictor of root growth rate for the different types of roots. In the rhizotrons, the root growth of lateral roots was found to be well correlated to the product of Da and the growth rate of the bearing root. Evaluation in field conditions from static observations attested consistent relationships between measured and predicted root length for lateral roots (slopes close to 1:1). The apical diameter can be considered as a good indicator of root growth potential, while actual lateral root growth depends on the bearing root elongation rate. Morphological static indicators calibrated from growth dynamics in rhizotrons are of major interest in explaining growth variability in field conditions. Especially the `growth rate-LAUZ' relationship can be considered a useful tool in interpreting field patterns of growing roots in relation to various soil conditions.     

杨树细根及草根的生产力与周转的研究

对北方杨树人工林0－40cm土层中杨树细根和草根（≤2mm）年生物量、分解量、死亡量、生长量和周转率进行观察研究。结果表明,杨树细根的年生物量为2．062t.hm^-2,死根生物量为0．746t.hm^-2,分解量为0．158t.hm^2,生长量为2．351t.hm^－2,周转率为每年1．14次,活草根的年生物量、死根生物量、分解量、生长量和周转率分别0．501、0．035、0．023、0．691t.hm^-2和1．38,同时给出了杨树细根干重损失随分解时间变化的方程：1nx/x0=0.9515e^-0.0014t。     

Effect of phosphorus deficiency on growth angle of basal roots in Phaseolus vulgaris

Root architectural plasticity might be an important factor in the acquisition by plants of immobile nutrients such as phosphorus (P). In this study, we examined the effect of P availability on the orientation of basal roots with respect to gravity, and thereby on the growth angle of these roots of common bean (Phaseolus vulgaris L.). In one set of studies the growth angle of basal roots of bean seedlings was measured over time. Sixteen bean genotypes were examined; six showed a decrease in root orientation with respect to gravity in low P media, one increased orientation, and nine showed no difference within 5 d of basal root emergence. Bean taproots also showed decreased root orientation with respect to gravity in low P. Growth angle after 5 d was correlated with field performance of contrasting genotypes in low P tropical soils. Mineral deficiencies other than P did not cause changes in root angle. In a split pouch system that provided high or low P solution to different parts of the root system, the decrease in root angle in low P was found to be a response to global P availability, and not local to the portion of the root system in low P. Effects of P availability on root angle were associated with reduced shoot P concentration, but preceded effects on plant biomass accumulation and leaf area expansion. Results from growth pouches for genotype G 19833 were confirmed using a solid-phase buffered sand-culture system supplying P at three levels. Pea (Pisum sativum), soybean (Glycine max Williams), chickpea (Cicer arietinum), lima bean (Phaseolus lunatus), and lentil (Lens culinaris) were grown with and without P; soybean and pea also showed decreased basal root angles in low P.     

Comparison of leaf life span, photosynthesis and defensive traits across seven species of deciduous broad-leaf tree seedlings

BACKGROUND AND AIMS: Leaf life span, photosynthetic parameters and defensive traits were compared across seven species of deciduous broad-leaved tree seedlings native to northern Japan to test the "cost-benefit hypothesis" that more productive leaves are more susceptible to herbivore attack than less productive leaves. METHODS: Studies were made on three early successional species, Alnus hirsuta, Betula maximowicziana and Betula platyphylla "japonica"; one mid-successional species, Ostrya japonica, and three late-successional species, Carpinus cordata, Quercus mongolica 'grosseserrata' and Acer mono. Photosynthetic parameters and defensive traits (total phenolics, condensed tannin and toughness) of leaves were measured for each species, and a bioassay test with Eri silkmoth larvae (Samia cynthia ricini) was undertaken to evaluate differences between species in susceptibility to herbivore attack. KEY RESULTS: Early successional species have a shorter leaf life span (62-88 d) than late successional species (155-187 d). Leaf nitrogen content and light-saturated photosynthetic rate per unit leaf area (P(sat)-area) and per unit leaf mass (P(sat)-mass) were negatively correlated with leaf life span. The nitrogen content of early successional species was about 30 mg g(-1) and that of late successional species was about 16 mg g(-1). Leaf toughness and the C/N ratio were positively correlated with leaf life span, although condensed tannin was not correlated with leaf life span. The bioassay test showed that the number of days the larvae survived was negatively correlated with leaf life span. Average survival of larvae feeding on leaves of A. hirsuta, which has the shortest leaf life span, was 14.4 d and that of Q. mongolica, which has the longest leaf life span, was 6.6 d. The number of days of larval survival was positively correlated with leaf nitrogen content. There was no correlation between days of larval survival and defensive traits. CONCLUSIONS: These results indicate that species with a shorter leaf life span have higher photosynthetic productivity and are more susceptible to herbivore attack than species with a longer leaf life span. This supports the "cost-benefit hypothesis".     

Modelling kiwifruit budbreak as a function of temperature and bud interactions

This paper presents two models of budbreak on canes of 'Hayward' kiwifruit (Actinidia deliciosa). A conventional 'chill unit' (CU) type model is compared with an alternative 'loss of potential' (LOP) approach, which assumes that the number of buds developing in spring depends on climate and node position-dependent bud-to-bud interactions that vary in duration and intensity. Both models describe how temperature, and application of a dormancy-breaking chemical, determine the overall amount of budbreak for whole canes. However, the LOP model does so by describing patterns of budbreak along canes. To do this, the cumulative influence of distal neighbours is assumed to cause a progressive fall in the capacity for bud development over the autumn-winter period, an influence that gets stronger as temperature rises. The LOP model also assumes that the rate of decline varies along the cane, as a function of some inherent bud property. These two factors mean that buds towards the base of the cane break less often under the suppressive influence of distal neighbours, while low temperature ('chilling') increases budbreak by diminishing the intensity of suppression relative to bud development rate. Under this scenario, dormancy-breaking chemicals (such as hydrogen cyanamide, HC) enhance budbreak by diminishing the duration of suppression. Models were calibrated using daily temperature series and budbreak proportion data from a multi-year regional survey, and were then tested against independent data sets. Both models were run from a fixed start date until the time budbreak was almost complete, or until a standard date. The fitted models described 87 % of variation in amount of budbreak due to site, year, HC and node position effects in the original data set. Results suggest that the correlation between chilling and the amount of budbreak can be interpreted as a population-based phenomenon based on interaction among buds.     

The position and growth of lateral roots on cultured root axes of tomato,Lycopersicon esculentum (Solanaceae)

Root axes of tomato (Lycopersicon esculentum) were cultured in vitro in three different concentrations of sucrose in order to vary their growth rate. Lateral root growth and the initiation of lateral root primordia were studied on each group of axes. Various aspects of primordium initiation, positioning, and emergence were quantified with a view to discovering variable and constant features of these processes. Variable parameters were the rate and frequency of root primordium emergence. Constant parameters, at least under the prevailing conditions, were the spacing between successive laterals and primordia, and the position of the primordia in relation to the vascular system. A model of primordium initiation is presented which combines controls determined by the divisional history of the potential primordium cell and by the vascular pattern.Dedicated with great respect to Prof. DrElisabeth Tschermak-Woess on the occasion of her 70th birthday in recognition of her distinguished contributions to cytology.     

Interactions between ethylene, gibberellin and abscisic acid regulate emergence and growth rate of adventitious roots in deepwater rice

Growth of adventitious roots is induced in deepwater rice (Oryza sativa L.) when plants become submerged. Ethylene which accumulates in flooded plant parts is responsible for root growth induction. Gibberellin (GA) is ineffective on its own but acts in a synergistic manner together with ethylene to promote the number of penetrating roots and the growth rate of emerged roots. Studies with the GA biosynthesis inhibitor paclobutrazol revealed that root emergence was dependent on GA activity. Abscisic acid (ABA) acted as a competitive inhibitor of GA activity. Root growth rate on the other hand was dependent on GA concentration and ABA acted as a potent inhibitor possibly of GA but also of ethylene signaling. The results indicated that root emergence and elongation are distinct phases of adventitious root growth that are regulated through different networking between ethylene, GA and ABA signaling pathways. Adventitious root emergence must be coordinated with programmed death of epidermal cells which cover root primordia. Epidermal cell death is also controlled by ethylene, GA and ABA albeit with cell-type specific cross-talk. Different interactions between the same hormones may be a means to ensure proper timing of cell death and root emergence and to adjust the growth rate of emerged adventitious roots.     

杉木,火力楠纯林及混交林细根周转的研究

系统研究了杉木、火力楠纯林及混交林细根生物量、生产力及年周转率。结果表明,杉木、火力楠纯林及混交林活细根生物量分别为８８０、３０３５和１５６０ｋｇ．ｈａ＾－１;死细根生物量为２３５、３９８和５６５ｋｇ．ｈａ＾－１;细根年周转率为１．２９、１．４２和１．４０;年生产量为１１３７、４３１８和２１７９ｋｇ．ｈａ＾－１;年死亡量为４９７．５９５和１１４９ｋｇ．ｈａ＾－１,分别相当于林分枯枝落叶年凋落量为３