A Developmental Pattern of Flowering in Colored <Emphasis Type="Italic">Zantedeschia</Emphasis> spp.: Effects of Bud Position and Gibberellin

The restricted flowering of colored cultivars ofZantedeschia is a consequence of developmental constraints imposed by apical dominance of the primary bud on secondary buds in the tuber, and by the sympodial growth of individual shoots. GA₃ enhances flowering inZantedeschia by increasing the number of flowering shoots per tuber and inflorescences per shoot. The effects of gibberellin on the pattern of flowering and on the developmental fate of differentiated inflorescences along the tuber axis and individual shoot axes were studied in GA₃ and Uniconazole-treated tubers. Inflorescence primordia and fully developed (emerged) floral stems produced during tuber storage and the plant growth period were recorded. Days to flowering, percent of flowering shoots and floral stem length decreased basipetally along the shoot and tuber axes. GA₃ prolonged the flowering period and increased both the number of flowering shoots per tuber and the differentiated inflorescences per shoot. Activated buds were GA₃ responsive regardless of meristem size or age. Uniconazole did not inhibit inflorescence differentiation but inhibited floral stem elongation. The results suggest that GA₃ has a dual action in the flowering process: induction of inflorescence differentiation and promotion of floral stem elongation. The flowering pattern could be a result of a gradient in the distribution of endogenous factors involved in inflorescence differentialtion (possibly GAs) and in floral stem growth. This gradient along the tuber and shoot axes is probably controlled by apical dominance of the primary bud. Online publication: 7 April 2005     

An appraisal of ear, leaf and stem 14CO2 assimilation, 14C-assimilate distribution and growth in a reproductive seed crop of amenity Lolium perenne

In seed crops of Lolium perenne, yield may be reduced by competition for a limited assimilate supply from sinks other than the ear. This study was undertaken to evaluate the priorities for allocation of assimilate within the crop from all photosynthetic sites on the main reproductive tiller after anthesis. Ear, stem and leaves were fed with ¹⁴CO₂ on two occasions; the assimilatory efficiency of these sources and the magnitude and pattern of ¹⁴C-assimilate export from each was determined. The growth of each part of the main tiller and subtending tillers was also measured. Stem elongation apparently dominated the current assimilate resource and the ear did not become a net importer of assimilate until this process had ceased. Assimilate allocation to the tillers was high throughout. The nature of any competition between these sinks is discussed. When crop growth was contrasted with that in a previous year, environmental factors were implicated as determinants of priority for assimilate allocation to each sink. Sources of carbon for seed filling are also discussed as is the relevance of these findings to seed crop management.     

Insights on the role of tillering in salt tolerance of spring wheat from detillering

Tillering is reduced by salinity, with the primary and secondary tillers being more affected than is the mainstem. To understand the importance of tillering in the salt tolerance of wheat plants, two contrasting genotypes of spring wheat (Triticum aestivum L.) were grown in a greenhouse under saline or non-saline conditions and were subjected to five progressive levels of detillering. Regardless of the genotype and salinity, shoot dry weight, seed yield and seed number per plant were all significantly decreased in the treatments where only one or two tillers per plant remained compared with the untouched treatment (more than three tillers), whereas these same variables per tiller tended to be increased on a per tiller (mainstem or substem tiller) basis. The increased seed yield per tiller observed with tiller reduction may be attributed to the enhanced seed number within the spikelet. Under saline conditions, the reductions in shoot dry weight, seed yield and seed number per plant for the salt-tolerant genotype Kharchia were of a greater magnitude in the treatments where only one or two tillers per plant were present compared with the untouched treatment, whereas the magnitude of this reduction in the salt-sensitive genotype Sakha 61 was decreased.     

GROWTH AND NUTRITION OF TIMOTHY (PHLEUM PRATENSE)

All tillers of timothy ( Phleum pratense ) growing in sand culture were labelled at weekly intervals. At each of eighteen dates between June and December a sample of plants was harvested, and dry weight and leaf area were determined for each group of tillers.
Tiller production was continuous throughout the experiment. Only those tillers which appeared before the end of July were able to form inflorescences, but among them the proportion of tillers with ears decreased with increasing lateness of origin.
Dry weight per plant increased rapidly for the first 14 weeks, and then did not change significantly until the end of the experiment. The flowering tillers lost dry weight after seed maturation; vegetative tillers, although smaller in size, continued to increase in number and weight. Dry weight per inflorescences was greatest in the main stem and decreased in subsequent tillers.
Leaf area per plant rose to a maximum soon after ear emergence and then declined, largely because the loss in leaf surface sustained by the flowering tillers was not offset by a corresponding increase in the vegetative tillers. Leaf area ratio decreased at first rapidly but later more gradually, while net assimilation rate was relatively high until maximum leaf area was almost attained, but then fell continuously to negligible values by October. Relative growth rate decreased throughout the experiment.
Initially the growth of the plant was largely determined by the flowering tillers in which, in close resemblance to the cereal plant, leaf area ratio and relative growth rate declined continuously. Tillers which did not flower were successively smaller in size and, because of their later origin, were exposed to less favourable environmental conditions by the time they had increased sufficiently in number to affect the growth of the plant.     

The patterns of growth, assimilation of 14CO2 and distribution of 14C-assimilate within vegetative plants of Loliumperenne at low and high density

The patterns of growth, assimilation of ¹⁴CO₂ and distribution of ¹⁴C-labelled assimilate were followed for 12 wk from sowing in individual plants of Lolium perenne grown in miniswards at either low (500 plants m^-2) or high (5000 plants m^-2) density. At the latter density, plants were characterised by a 50% reduction in RGR, by the production of fewer tillers, especially second- and third-order tillers, and by a reduction in mean tiller weight. All the green and senescing leaves of each tiller assimilated ¹⁴CO₂ and the overall assimilatory capacity of a tiller was directly related to its dry weight. At both densities the plant consisted of a main shoot and established tillers with comparable assimilatory activities and a range of developing tillers that assimilated relatively small amounts of ¹⁴CO₂. As each successive primary tiller developed it was supplied with assimilate from the main shoot and the degree of support was inversely proportional to the dry weight of the tiller. At both densities it was concluded that the first primary tiller could be regarded as an independent assimilatory unit when it reached a dry weight of about 25 mg even though some import of main shoot assimilate continued until the tiller was double this weight. The supply of assimilate to the root system was greatly reduced at both densities compared with previous observations on plants grown singly.     

不同穗型小麦品种分蘖发育的代谢基础研究

比较研究了大穗型和多穗型小麦品种的分薛发育特征及其代谢基础。结果表明,与多穗型品种相比,单株分蘖较少的大穗型品种分蘖期具有较低的IAA氧化酶（IAAO）活性、较高的IAA含量和较强的碳代谢活性。在分蘖的两极分化期,大穗型品种在碳、氮代谢和同化物分配方面存在较强的主茎优势,因此,其分蘖较难继续发育成穗。     

GROWTH AND NUTRITION OF TIMOTHY (PHLEUM PRATENSE)

Individual tillers of timothy ( Phleum pratense ) growing from seed in pots were labelled and their life history studied over a period of two years.
The leaf axil position of the first-produced tiller depended on its time of origin. A minimum of five leaves on the parent axis was necessary before its appearance, and no tiller emerged from the axil of any of the three terminal leaves of a shoot.
Ear-bearing capacity was high in early-formed tillers, but decreased progressively with later dates of origin despite favourable day length; all tillers arising after the end of July failed to produce inflorescences. A greater proportion of tillers on the main stem formed ears than secondary tillers of the same age. Time of ear emergence varied only within a few days for all tillers arising before the end of April. The number of leaves formed up to ear emergence varied linearly with date of origin and tiller position, the lowest recorded number being six.
Dry weight and length of inflorescence as well as seed yield per tiller varied linearly with the date of origin of tillers. Primary tillers generally had longer and heavier inflorescences and produced more seed per stem than secondary tillers of the same age.
Tillers arising at all times were found to have a limited span of life, not normally exceeding just over a year. The mean age of ear-bearing tillers varied linearly with their date of origin. Death-rates of all tillers generally increased with their age. Relative death-rates of fertile and vegetative tillers rose rapidly in the autumn but decreased during winter and early spring. Vegetative tillers which elongated in the year of formation survived for a shorter time than equivalent tillers in which stem elongation was delayed until the following year.
The perennial habit of grasses is discussed in relation to tiller survival. Successive new formation and death of tillers keep the plant in a condition of continuous dynamic change.     

Regulation of reproduction in wild and cultivatedLycopersicon esculentum Mill. by vesicular-arbuscular mycorrhizal infection

Summary We examined how mycorrhizal infection byGlomus etunicatum Becker and Gerd. affected flowering phenology and components of reproduction in eight wild accessions and two cultivars ofLycopersicon esculentum Mill. We did this by performing a detailed demographic study of flower, fruit and seed production. Mycorrhizal infection had variable effects on the ten accessions. Infection significantly decreased the time taken to initiate flowering in some accessions. In addition, infection increased flowering duration in some accessions. In many accessions, infection significantly increased seed production, primarily by increasing the number of inflorescences and infructescences. In some accessions, mycorrhizal infection also increased the proportion of flowers that produced mature fruits or the number of seeds per fruit. Among accessions, shoot phosphorus content was correlated with seed productivity for both mycorrhizal and non-mycorrhizal plants. However, non-mycorrhizal plants produced more seed biomass per mg of shoot phosphorus than mycorrhizal plants.     

A Quantitative Analysis of the Uptake of Carbon and of the Supply of 14C-labelled Assimilates to Areas of Meristematic Growth in Lolium temulentum

A quantitative analysis of the ¹⁴C-labelled assimilate suppliedby leaves on the main shoot to terminal meristem, stem, tillers,and roots was conducted during parallel periods of reproductiveand vegetative development in Lolium temulentum. The initial rate of entry of carbon into the shoot varied withthe area and photosynthetic efficiency of the assimilating leaf.Subsequently, respiratory losses of carbon during translocationand incorporation of assimilate at the site of utilization alsovaried. The combined effect of these differences resulted inthe supply of recently assimilated carbon being twofold greaterin reproductive shoots than in vegetative shoots, while withinshoots the carbon supply of the youngest fully expanded leafranged from four-or five-fold greater than the oldest leaf inyoung shoots, to two-or three-fold greater in older shoots.In both reproductive and vegetative shoots, the two or threeyoungest leaves thus dominated the supply of carbon for meristematicgrowth. Meristematic tissue in expanding leaves and leaf primordia atthe terminal meristem of the vegetative shoot received 18–27per cent of the total shoot carbon. This meristem utilized aboutthe same proportion of shoot carbon when it developed into aninflorescence, indicating no major change in the level of meristematicactivity. The proportion of shoot carbon utilized in stem growthincreased as both reproductive and vegetative shoots aged; thisincreased meristematic activity in stem internodes was accompaniedby reduced export of carbon to roots, which received less than10 per cent of the shoot carbon when the experiments ended.The main shoot translocated 20–30 per cent of its recentlyassimilated carbon to developing and rooted tillers, which assinks for carbon were thus as important as the terminal meristemand stem. This outward flow of carbon continued relatively uncheckedwhen donor and receptor shoots developed inflorescences.     

Effects on the Stem of Winter Barley of Manipulating the Source and Sink during Grain-filling: I. CHANGES IN ACCUMULATION AND LOSS OF MASS FROM INTERNODES

Source and sink activities of field-grown barley plants weremanipulated by shading the whole plant or by de-earing at varioustimes after anthesis. The changes in the masses of the internodes,ears, grains and leaves were followed until the grain matured. Shading reduced the mass of ears and grains and, in addition,shading to 45% of incident radiation from anthesis preventedmature internodes from increasing in mass. Severe shading (to9% of incident radiation) from 11 d after anthesis caused thenormal decrease in mass of internodes to occur earlier thanin unshaded plants. Thus, only assimilate in excess of thatrequired for tructural growth was stored in internodes. Lowerinternodes of plants shaded from anthesis lost more of the massthat had been accumulated prior to anthesis than control plants,giving a greater potential pre-anthesis contribution to grainyield. Internodes from plants that were de-eared accumulated more massthan internodes of control plants in one year but not in another,i.e. the potential for internodes to store assimilate appearsto be exploited in some years but not others. The mass of internodesfrom de-eared plants either decreased after that of controlplants or their mass stayed constant. De-eared plants had anincreased number of tillers compared with control plants; thesetillers could have been an alternative sink for assimilate storedin the internodes of de-eared plants. Mobilization of assimilate from internodes is in response todemand for assimilate from grains being in excess of currentphotosynthesis. Key words: Winter barley, stem reserves, mobilization, shading, de-earing     

Resource availability regulates reproductive meristem activity, development of reproductive structures and seed set in buckwheat (Fagopyrum esculentum)

Grain yields in buckwheat ( Fagopyrum esculentum ) are consistently low despite a profuse and long-lasting flowering. The mechanisms underlying this reproductive failure are still largely unknown. In this study, performed in strictly controlled conditions, the possible implication of assimilate availability in the regulation of the reproductive development was investigated in the 'La Harpe' cultivar, by manipulating sink/source relationships through excisions of selected leaves and/or selected inflorescences and hand pollinations of selected and precisely identified flowers. Increasing or decreasing source/sink organ ratio in buckwheat, respectively, enhanced or reduced the production of racemes by the shoot apical meristem and of cymes by the racemes, ultimately modulating the number of flowers per plant. The effect on seed production was essentially indirect and related to flower production. Seeds developing on a plant also affected reproductive morphogenesis, limiting flower production. Three stages at which flower development to mature seed may fail have been identified. First, an abnormal morphogenesis resulted in flowers with an undersized and sterile gynoecium. Secondly, after anthesis, numerous flowers with a normal sized gynoecium, were unable to develop a seed after hand pollination. Finally, a small proportion of flowers exhibited a later abortion of the seed. Flower fate is dependent on position in the raceme and time of opening: chance to develop normally and produce a ripe seed is higher for flowers at the base than at the top of the inflorescence. A mechanism, internal to the raceme, regulates flower fate, independently of assimilate availability. This mechanism may be disturbed by drastic reductions in nutrient supply.     

Additive and non-additive effects of simulated leaf and inflorescence damage on survival, growth and reproduction of the perennial herb Arabidopsis lyrata

Herbivores may damage both leaves and reproductive structures, and although such combined damage may affect plant fitness non-additively, this has received little attention. We conducted a 2-year field experiment with a factorial design to examine the effects of simulated leaf (0, 12.5, 25, or 50% of leaf area removed) and inflorescence damage (0 vs. 50% of inflorescences removed) on survival, growth and reproduction in the perennial herb Arabidopsis lyrata. Leaf and inflorescence damage negatively and independently reduced flower, fruit and seed production in the year of damage; leaf damage also reduced rosette size by the end of the first season and flower production in the second year. Leaf damage alone reduced the proportion of flowers forming a fruit and fruit production per plant the second year, but when combined with inflorescence damage no such effect was observed (significant leaf × inflorescence damage interaction). Damage to leaves (sources) caused a greater reduction in future reproduction than did simultaneous damage to leaves and inflorescences (sinks). This demonstrates that a full understanding of the effects of herbivore damage on plant fitness requires that consequences of damage to vegetative and reproductive structures are evaluated over more than 1 year and that non-additive effects are considered.     

The effect of tiller removal and tiller defoliation on competition between the main shoot and tillers of spring barley

In glasshouse and field experiments the source-sink relations of the main shoot of plants of spring barley were modified by tiller removal and tiller defoliation. Decreasing competition by tiller removal promoted the growth of the residual main shoot and its component parts, and the earlier tillers were removed the greater was the effect. Stem dry weight was increased four-fold in the glasshouse by early tiller removal and was doubled in the field experiment. The grain yield of the main shoot ear was increased by 26 – 30% by tiller removal compared with tillering control plants and this was due to larger grains in all spikelet positions. On the other hand increasing competition by regular tiller defoliation had relatively little effect on the growth and development of the main shoot in the glasshouse study, but in the field the main shoot grain yield was reduced by 10% compared with the control. The main effect of tiller defoliation was on the development of tillers. In the glasshouse tillers survived repeated defoliation, continued to be produced, and the majority produced grain but with fewer and smaller grains per ear than in control plants. Tiller growth was supported by the import of assimilate from the main shoot and this was accompanied by an increase in the photosynthetic rate of the main shoot leaves. In the field all defoliated tillers died within 4 wk. These responses are discussed in terms of the physiological interrelations between the main shoot and tillers.     

Seasonal variation in honeyeater foraging behaviour,infloresence abundance and fruit set in Banksia spinulosa (Proteaceae)

Seasonal variation in the foraging behaviour of honeyeaters and the production of fruit were examined in relation to the flowering intensity of Banksia spinulosa over two flowering seasons. The abundance of inflorescences was greater in the mid than in the early and late periods of the flowering season. In the mid period, many plants were blooming and each plant had many flowering inflorescences. Inflorescences received most visits by honeyeaters in the early flowering period; the visitation rate declining as flowering progressed. Eastern spinebills were the most common floral visitors at all times during the season. The number of foraging probes made at inflorescences by eastern spinebills did not differ throughout the season. Foraging movements between inflorescences on the same plant were more frequent in the mid period than in the early and late periods. Long distance movements between plants (more than 10 m apart) were promoted by aggressive interactions between honeyeaters. Inflorescences flowering in the late period were less likely to develop follicles because there were fewer visits by birds and/or because resources had been allocated to inflorescences pollinated earlier in the season. The number of follicles produced per infructescence did not differ between flowering periods. Overall, the number of inflorescences produced per plant, the number of visits received per inflorescence and the proportion of inflorescences that developed follicles were greater in 1987 than in 1988.     

Factors affecting fruit and seed production of Paeonia ostii “Feng Dan”, an economically important oil tree

Many factors may affect reproduction of animal-pollinated species. In this study, the effects of pollen limitation, attractive traits (flower number, plant height and flower width) and flowering phenological traits (flowering onset, duration and synchrony) on female reproduction, as well as the patterns of variation in fruit and seed production within plants, were investigated in Paeonia ostii “Feng Dan” over two flowering seasons (2018 and 2019). Fruit set was very high (90%), and pollen supplementation did not increase fruit and seed production in either year, indicating no pollen limitation. Fruit set, ovule number per fruit and mean individual seed weight per fruit were not affected by any of the six attractive and phenological traits in either year, whereas seed number per fruit was related to the three attractive traits in one or both years. Seed number per plant was positively affected by the three attractive traits and best explained by flower number in both years, but the effect of each of the three phenological traits on seed number per plant differed between years. Within plants, the fruit set, ovule number, seed set and seed number per fruit declined from early- to late-opening flowers, presumably because of resource preemption, but the mean individual seed weight did not vary across the flowering sequence. Our study shows that attractive traits of Paeonia ostii “Feng Dan” are more important than flowering phenological traits in the prediction of total seed production per plant.     

The effect of light environment, leaf area, and stored carbohydrates on inflorescence production by a rain forest understory palm

Variation in flowering by long-lived plants may be correlated with current resource availability. If, however, there are trade-offs between current and future reproduction, or between reproduction and storage or growth, then understanding variation requires a whole-plant, longer-term perspective. Inflorescence production by Calyptrogyne ghiesbreghtiana Linden ex. H. Wendl., an understory palm, was studied over 3 years. Annual inflorescence production varied greatly and was correlated with variation in plant size and light environment. There was no trade-off between past inflorescence production and the frequency of future inflorescence production. On the contrary, individuals that produced more inflorescences than predicted from their size and light environment tended to continue to do so in subsequent years also. I manipulated the resource environment of a subset of plants by removal of leaves and/or reproductive spikes. Leaf removal suppressed inflorescence production for the following 2 years, but spike removal had no effect. One year after leaf removal stored reserves were, on average, back to pre-treatment levels. There was, however, a negative effect of recent inflorescence production on storage. Plants with higher levels of storage had higher inflorescence production in the next 75 days. In C. ghiesbreghtiana the resource cost of reproduction is apparent in short-term variation in stored reserves. In contrast, annual inflorescence production does not follow a trade-off pattern between successive years, but consistently reflects both plant size and the light environment. Received: 20 October 1996 / Accepted: 25 January 1997     

Tillers Do Not Influence Phytotoxicity of Imazamethabenz in Wild Oat (Avena fatua)

The response of wild oat to imazamethabenz varies with the growth stage, but the role of tillers in this regard is unclear. Removal of tillers at the three-leaf stage before spraying with imazamethabenz did not significantly affect the total shoot fresh weight measured 3 weeks later. The leaf area and dry weight of intact plants at the three-leaf stage were 17–21% greater than for plants with coleoptilar and first leaf main shoot tillers (T0 and T1) removed. The greater leaf area may have increased herbicide interception per plant. Similar fresh weight reductions in main shoot, total tillers, and total shoots were found whether imazamethabenz was applied to the plant at the two-leaf without tillers or the three-leaf with two tillers stage. Imazamethabenz applied only to the main shoot reduced total shoot dry weight more than an equivalent amount of imazamethabenz applied only to tiller T1 or applied over the whole shoot. Imazamethabenz had the least inhibitory effect on whole plant growth when applied only to T1. When ¹⁴C-herbicide was applied to the first main shoot leaf of plants at the three-leaf stage with two tillers, the ¹⁴C translocated 38% to roots, 33% to the main shoot, and nearly 30% to all tillers. When ¹⁴C-herbicide was applied to the first leaf of T1 then the ¹⁴C translocated 50% to T1, 25% to the main shoot, 20% to roots, and 5% to all other tillers. The translocation pattern and fresh weight values suggested that the presence of early tillers during herbicide application neither increased nor decreased imazamethabenz efficacy in wild oat. Received June 4, 1997; accepted June 5, 1997     

The Distribution of Assimilates in Lolium multiflorum Lam. Following Differential Defoliation

In seedling plants of Lolium multiflorum Lam. the tillers weredefoliated but the main shoot was left intact. Radiocarbon as¹⁴CO₂ was supplied to this shoot at different times followingtiller defoliation and the pattern of distribution of labelledassimilates was determined quantitatively. It was found thata greater proportion (approximately 10–20 per cent) ofexported assimilate was translocated to the cut tillers butalthough the proportion supplied to the root system was lessthe total radiocarbon incorporated by the roots was unchanged.This was brought about by a large increase in the export ofradiocarbon fixed by the intact shoot—up to 100 per centfollowing one treatment. These alterations in the organizationof the defoliated plant lead to a greater efficiency in thecarbon economy and are discussed in relation to the stress imposedby defoliation.     

Seed production of cut-leaved teasel (<Emphasis Type="Italic">Dipsacus laciniatus</Emphasis>) in central Missouri

Cut-leaved teasel is an invasive weed in Missouri that reduces the diversification of native species along roadsides and impairs traffic visibility. Teasel is a biennial and grows as a rosette in the first year and flowers the second year. Reproduction is only by seed. Field studies were conducted in 2004 and 2005 at two locations to assess the seed production of cut-leaved teasel. From a natural stand, fifteen plants were tagged at the onset of flowering. Selected plants included those considered growing in a group and those growing alone; a plant was considered alone when no other plant was adjacent for at least 60 cm. Whenever a seedhead completed flowering, it was covered with a cellophane bag and harvested one month later. Linear regression was used to correlate the weight of seeds from a single seedhead and number of seeds to estimate the total seed production per seedhead. The number of seedheads per plant varied from 3 to 56. On average, plants growing alone had 64% more seedheads per plant than plants occurring in a group. Seed numbers in the primary seedhead ranged from 511 to 1,487. Total seed production per plant ranged from 1,309 to 33,527. Seed production was 61% greater for plants growing alone versus those growing in a group and was more prolific in 2005 than in 2004. In addition, seed production per plant varied between locations for plants growing alone, but seed yield per plant was similar for plants growing in groups. Colonization of teasel in new areas is facilitated by higher seedhead numbers per plant and total seed production compared to reproduction of plants in areas of intraspecific competition.     

Root-Tiller Interrelationships in Spring Barley (Hordeum distichum (L.) Lam.)

Root-tiller relations were investigated in spring barley grownin soil in deep pots. The total dry wt of the root system reachedits maximum 6 weeks from sowing, when the shoot weight was only50 per cent of its value at maturity. Seminal and nodal rootscomprised 40 and 60 per cent, respectively, of the total rootdry wt at maturity; the majority of the nodal root weight wasassociated with the main shoot. The main shoot had approximatelytwice as many nodal roots as either of the first two primarytillers (T1 and T2), and the primary and secondary tillers appearinglater were very poorly rooted. Some tillers, especially secondarytillers that died prematurely, produced no nodal roots. Theweight of the seminal roots and nodal roots attached to themain shoot continued to increase up to maturity but the drywt of nodal roots on tillers declined with time. This patternof growth was closely related to the pattern of ¹⁴C assimilateddistribution within the root system. A very small proportionof ¹⁴C assimilated by the main shoot and T1 and T2 was exported.The majority of the exported assimilate went to the seminalroot system and to nodal roots attached to the main shoot. Individualnodal and seminal roots seemed to have different roles in supplyingnutrients to the shoot system, with the former mainly providing³²P-phosphate to its tiller of origin and the latter generallysupplying the main shoot and primary tillers. Hordeum distichum. (L.) Lam., barley, root growth, nodal roots, seminal roots, tillering, assimilate distribution, ³²P-distribution