Understanding the germination of bulbils from an ecological perspective: a case study on Chinese yam (Dioscorea polystachya)

Background and Aims

Bulbils serve as a means of vegetative reproduction and of dispersal for many plants; this latter aspect making them analogous to seeds. However, germination of bulbils may differ considerably from seeds due to dissimilar anatomical structures and perhaps environmental cue perception. The few laboratory studies done on bulbils suggest that their germination is similar to that of seeds in the same habitats and to vegetative buds of winter-dormant plants. The present study is the first to examine how bulbil germination is controlled in nature in relation to dispersal (before vs. after winter of the same cohort) and to ambient temperatures.

Methods

Under laboratory conditions, temperature and light requirements for root and shoot emergences from bulbils of Dioscorea polystachya collected in September, 2005, February, 2006 (produced in 2005) and July, 2006 were determined. Effects of cold stratification and dry storage for releasing dormancy were tested on September and July bulbils. The phenology of dormancy release and of root and shoot emergences and the persistence of bulbils in soil were followed over time under field conditions.

Key Results

Although a low percentage of bulbils collected in July or in September produced roots, but no shoots, in the laboratory and field, these roots died within approx. 1 month. Regardless of collection date, cold stratification markedly increased root and shoot emergences. Bulbils sown outdoors in October produced roots and shoots the following March and April, respectively. The soil bulbil bank is short lived.

Conclusions

Bulbils of D. polystachya are similar to seeds of many temperate plants being mostly dormant when dispersed in summer or autumn and overcoming dormancy with cold stratification during winter. Adaptively, bulbil germination primarily occurs in spring at the beginning of a favourable period for survivorship and growth.     

Studies on the growth and development of Oxalis latifolia

The growth and development of Oxalis latifolia was assessed on outdoor-grown potted plants raised from bulbs. Three main phases of growth appeared to be correlated with major physiological developments in the aerial and subterranean organs. Establishment (weeks 1 – 5) was characterised by a slow gain in plant dry weight accompanied by high relative growth rate (RGR) values for individual plant organs. The second phase of growth (weeks 6 – 12) was reproductive, as stolons grew and initiated bulbils at the apices. Bulbils had the capacity to produce their own shoot system and ultimately some primary bulbils supported secondary bulbils. Dry matter accumulation was in the order: bulbils > peduncles > leaves > petioles > stolons = roots > parent bulb. The RGR of individual plant organs declined together with the net assimilation (NAR) rates and the leaf area ratio of the shoots during the reproductive phase and beyond into the senescence phase (week 13 onwards). The results are discussed in relation to the problem of achieving weed control via cultural and herbicide treatments.     

Biology of the polyploid geophyte Allium oleraceum (Amaryllidaceae): Variation in size,sexual and asexual reproduction and germination within and between tetra-, penta- and hexaploid cytotypes

It has been proposed that plant species cytotypes commonly exhibit altered morphology, reproduction, geographic and ecological distributions. We studied phenotypic variation in height, sexual (flowers, seeds) and asexual (aerial bulbils) reproduction in natural populations and in the conditions of a common garden of three cytotypes (2n = 4x, 5x, 6x) of the bulbous geophyte Allium oleraceum in the Czech Republic. Additionally, we compared the germination and dormancy of seeds and bulbils to determine whether propagules have different ecological roles. The pattern of morphological differentiation observed between cytotypes under natural conditions was similar to that observed under common garden conditions, suggesting that variability in morphological characteristics appears to be associated with ploidy levels. We revealed differences in size and sexual and asexual production between A. oleraceum cytotypes, but with wide overlap among cytotypes, suggesting a limited possibility of the studied traits to reliably distinguish between cytotypes. Tetraploids and pentaploids were rather similar; they were taller and produced more flowers than hexaploids, which were mostly flowerless (mean <0.7 flower/plant). All cytotypes were able to produce viable seeds, but their numbers were extremely low, usually less than 3 seeds per inflorescence; clonal reproduction via aerial bulbils dominates in all cytotypes (flower:bulbil ratio <0.5), with tetraploids producing more but lighter bulbils than other cytotypes. The seed: ovule ratio was low (<0.1) in all cytotypes, although hexaploids reached higher values than other cytotypes. Bulbils germinated better (means >80%) than seeds in all cytotypes, with pentaploid bulbils showing the highest germination (mean 90.5%). The cytotypes did not differ in seed germination (range of means 73.4%–76.3%). About 6% of seeds did not germinate but were still viable at the end of the experiment, while all non-germinated bulbils of all cytotypes had rotten away. Seeds, but not bulbils, can likely form a short-term persistent propagule soil bank. We found no evidence of a phenotypic trade-off between the production of flowers (seeds) and the production of bulbils within the inflorescence of all studied cytotypes. Together, inter-cytotypic differences in fitness-related traits cannot completely explain the different geographic and ecological distributions of cytotypes in the Czech Republic found by previous research.     

Biomass and production of fine and coarse roots during regrowth of a disturbed subtropical humid forest in north-east India

Seasonal variation and depthwise distribution of dry matter in roots of different diameter classes and their annual production were studied using sequential core sampling. The investigations were carried out in three stands of a subtropical humid forest of north-east India representing different stages of regrowth after tree cutting. The mean annual standing crop of fine (<2 mm in diameter) and coarse (2–15 mm diameter) roots increased gradually from 5.4 Mg ha^-1 and 0.7 Mg ha^-1 in 7-yr old regrowth to 9.4 Mg ha^-1 and 2.8 Mg ha^-1 in 16-yr old regrowth, respectively. The contribution of fine roots to the total root mass declined from 88% in 7-yr old regrowth to 77% in both 13 and 16-yr old regrowths, while that of coarse roots increased from 12 to 23%. A major portion of fine roots (59–62%) was present in 0–10 cm soil layer, but the coarse roots were concentrated in 10–20 cm soil depth (38–48%). In all the three stands, biomass of both fine and coarse roots followed a unimodal growth curve by showing a gradual increase from spring/pre-rainy season to autumn/post-rainy season. Biomass to necromass ratio increased from 2.5 in the 7-yr old to 3.2 in the 16-yr old stand. The annual fine root production increased from 5.9 Mg ha^-1 to 7.7 Mg ha^-1 and total root production from 7.6 Mg ha^-1 to 14.7 Mg ha^-1 from 7-yr to 16-yr old regrowth.     

Insurance against reproductive failure in a semelparous plant: bulbil formation in Agave macroacantha flowering stalks

Bulbils are small aerial rosettes that occur on the flowering stalks of semelparous Agave plants and in related families, and that are capable of acting as clones of the parent plant. We hypothesized that bulbil formation was inversely related to fruiting success in the flowering stalk, and we explored this hypothesis in A. macroacantha, a species from South-Central Mexico. Forty randomly chosen plants were divided amongst three treatments: (a) elimination of all floral buds, (b) exclusion of pollinators, and (c) control. We also studied 22 plants in which the flowering stalk had been felled by goat grazing. Between September and November 1991 we kept a record of the numbers of bulbils and capsules produced in each flowering stalk. Significant (P<0.0001) differences between treatments were found in the proportion of plants hearing capsules and bearing bulbils. The control treatment had the highest proportion of plants producing capsules, treatment a had the highest proportion of individuals bearing bulbils, while treatment b showed an intermediate response. In the goat-grazed group, 45% of the plants failed to produce any propagative structure after the stalk was cut, and half of all plants produced bulbils on the remaining stump. A significant inverse relationship between the numbers of capsules and the numbers of bulbils per plant was found for the three randomly assigned treatments. Our results suggest that once the production of the flowering stalk has been triggered and the death of the rosette is irreversible, bulbils may act as an insurance mechanism that increases the probability of successful reproduction of the genet.     

Germinating seeds or bulbils in 87 of 113 tested Arctic species indicate potential for ex situ seed bank storage

Arctic plant species are expected to lose range due to climate change. One approach to preserve the genetic and species diversity for the future is to store propagules in seed vaults. However, germinability of seeds is assumed to be low for Arctic species. We evaluated ex situ storage potential of 113 of the 161 native angiosperms of Svalbard by studying seed ripening and germination. Seeds or bulbils were collected, and germinability was tested after one winter of storage in the Svalbard Global Seed Vault. Twenty-six of the species did not produce ripe propagules, 8 produced bulbils, and 79 produced seeds. Bulbils sprouted to high percentages. Seeds of 10 species did not germinate, 22 had low germination (<20 %), 34 had germination of 21–70 %, and 13 had high germination percentages (>70 %). More than 70 % of the species belonging to Asteraceae, Brassicaceae, Caryophyllaceae, Juncaceae, Rosaceae, and Saxifragaceae germinated. Cold tolerant, common species had higher germination percentages than relatively thermophilous, rare species. Germination percentages were six times higher than observed in 1969 (n = 51) and 0.7 times that observed in 2008 (n = 22), indicating that recent climate warming improves germination in the Arctic. While in situ conservation is of vital importance, ex situ conservation in seed banks is a potential complementary conservation strategy for the majority of Arctic vascular plant species. For species that did not germinate, other methods for ex situ conservation should be sought, for example, growing in botanical gardens.     

Dormancy in bulbils of several herbaceous plants: Effects of photoperiod,light, temperature,oxygen and gibberellic acid

Effects of some environmental conditions (photoperiod, white and colored lights, temperature, partial oxygen pressure) and growth regulators (gibberellic acid, 2-chloroethyltrimethylammonium chloride) on induction and release of dormancy of the bulbils ofDioscorea batalas, Laportea bulbifera, Elatostema involucratum andSedum bulbiferum were investigated. Bulbils were formed under short-day conditions inLaportea andElatostema, under long-day conditions inSedum, and irrespective of photoperiods inDioscorea. In all species exceptSedum, immature bulbils required light, particularly blue or far red, for sprouting (photo-sprouting stage), and mature bulbils required a cold treatment (thermo-sprouting stage). The duration of photo-sprouting and thermo-sprouting stages and the degree of dependency on light or low temperature of sprouting differed from species to species. Sprouting of chilled mature bulbils of these species was promoted by light, especially by red or green light. Both immature and mature bulbils ofSedum sprouted under short-day conditions. Continuous irradiation with blue, far-red and green light markedly inhibited their sprouting. Oxygen at high concentration inhibited the sprouting of immature bulbils inDioscorea; in the other species it promoted sprouting regardless of the maturation of the bulbils. Applications of gibberellic acid caused the sprouting of bulbils the absence of light, chilling or photoperiodic treatment in all species exceptDioscorea, in which gibberellic acid inhibited sprouting. Polyphenol oxidase activity was very high in the homogenates ofDioscorea bulbils, and increased further when the bulbils had been treated with gibberellic acid. In the other species, little or no such activity was observed.     

卷丹珠芽萌发的组织学观察

以卷丹(Lilium laneifolium)珠芽为试材,采用野外调查法、石蜡切片法、徒手切片法、离析法,观察珠芽各部分形态和结构,用分光光度法测定各片鳞叶的花青素和光合色素含量,为其珠芽繁殖生物学研究提供资料。结果表明,珠芽由鳞叶、鳞茎和不定根构成,鳞叶外表皮细胞具有发达的角质层,外表皮内侧1～2层叶肉细胞含有花青素;叶肉细胞含绿色造粉体,第1～3片鳞叶基部的绿色造粉体向不定根伸长方向集中分布,鳞叶色素含量由外至内逐渐降低;鳞叶维管束为外韧维管束。鳞茎主要由皮层和维管柱构成,鳞茎上端包括顶端分生组织和芽鞘,在下端细胞部分发生程序性死亡,但未发现类似叶片脱落时叶柄基部出现的离层结构。不定根起源于第2片鳞叶基部环生的鳞茎皮层细胞,不定根与周围鳞叶组织分离。在珠芽萌发过程中鳞叶的物质供给出现分化现象。     

Does salinity reduce growth in maize root epidermal cells by inhibiting their capacity for cell wall acidification?

The reduction in growth of maize (Zea mays L.) seedling primary roots induced by salinization of the nutrient medium with 100 millimolar NaCl was accompanied by reductions in the length of the root tip elongation zone, the length of fully elongated epidermal cells, and the apparent rate of cell production: Each was partially restored when calcium levels in the salinized growth medium were increased from 0.5 to 10.0 millimolar. We investigated the possibility that the inhibition of elongation growth by salinity might be associated with an inhibition of cell wall acidification, such as that which occurs when root growth is inhibited by IAA. A qualitative assay of root surface acidification, using bromocresol purple pH indicator in agar, showed that salinized roots, with and without extra calcium, produced a zone of surface acidification which was similar to that produced by control roots. The zone of acidification began 1 to 2 millimeters behind the tip and coincided with the zone of cell elongation. The remainder of the root alkalinized its surface. Kinetics of surface acidification were assayed quantitatively by placing a flat tipped pH electrode in contact with the elongation zone. The pH at the epidermal surfaces of roots grown either with 100 millimolar NaCl (growth inhibitory), or with 10 millimolar calcium ± NaCl (little growth inhibition), declined from 6.0 to 5.1 over 30 minutes. We conclude that NaCl did not inhibit growth by reducing the capacity of epidermal cells to acidify their walls.     

Decomposition and nutrient release of grass and tree fine roots along an environmental gradient in southern Patagonia

Decomposition of fine roots is a fundamental ecosystem process that relates to carbon (C) and nutrient cycling in terrestrial ecosystems. However, this important ecosystem process has been hardly studied in Patagonian ecosystems. The aim of this work was to study root decomposition and nutrient release from fine roots of grasses and trees (Nothofagus antarctica) across a range of Patagonian ecosystems that included steppe, primary forest and silvopastoral forests. After 2.2 years of decomposition in the field all roots retained 70–90% of their original mass, and decomposition rates were 0.09 and 0.15 year^?1 for grass roots in steppe and primary forest, respectively. For N. antarctica roots, no significant differences were found in rates of decay between primary and silvopastoral forests (k = 0.07 year^?1). Possibly low temperatures of these southern sites restricted decomposition by microorganisms. Nutrient release differed between sites and root types. Across all ecosystem categories, nitrogen (N) retention in decomposing biomass followed the order: tree roots > roots of forest grasses > roots of steppe grasses. Phosphorus (P) was retained in grass roots in forest plots but was released during decomposition of tree and steppe grass roots. Calcium (Ca) dynamics also was different between root types, since trees showed retention during the initial phase, whereas grass roots showed a slow and consistent Ca release during decomposition. Potassium (K) was the only nutrient that was rapidly released from both grass and tree roots in both grasslands and woodlands. We found that silvopastoral use of N. antarctica forests does not affect grass or tree root decomposition and/or nutrient release, since no significant differences were found for any nutrient according to ecosystem type. Information about tree and grass root decomposition found in this work could be useful to understand C and nutrient cycling in these southern ecosystems, which are characterized by extreme climatic conditions.     

Quantitative changes of batatasins and abscisic acid in relation to the development of dormancy in yam bulbils

As yam bulbils (Dioscorea batatas Decne.) grow on the motherplants, dormancy of the bulbils develops gradually. This isevidenced by the fact that the sproutability of bulbils decreasedin proportion to their sizes, 2–4 mm to 8–12 mmin diameter when bulbils are detached and sown in Petri dishes.This decreasing sproutability was seen until the stratificationat 5°C for dormancy release was completed. During the growth process of attached bulbils, the contentsof batatasins I and II increased markedly per surface area andper bulbil although on a fresh weight basis they remained constant.The increases of batatasin III and abscisic acid (ABA) weremore striking and were evident in all comparisons. These results corresponded to different localization of batatasinsand ABA in bulbil issues. All of 3 batatasins were localizedin the outermost layers of the bulbils, whereas ABA was distributedevenly throughout their tissues. These results suggest differentroles for batatasins and ABA in the regulation of yam bulbildormancy. ¹ This paper concerns K. H.'s Ph. D. dissertation presentedto Tohoku University based on work done at I. P. C. R. ² Present address: Biological Institute, Faculty of Science,Tohoku University, Sendai, Japan. (Received November 15, 1972; )     

Floral structure and ontogeny in Globba (Zingiberaceae)

We present new comparative morphological and ontogenetic data on flowers and bulbils of Globba (Zingiberaceae) to clarify their homologies. Globba flowers are characteristically Zingiberaceous, possessing a single stamen and epigynous (``supragynopleural') nectaries, but are unusual as the anther bears triangular lateral outgrowths and the style is held tightly in position across the curvature of the filament like a bowstring. Floral ontogeny in Globba is similar to other Zingiberaceae. Characteristic features, such as anther wings, occur late in development, shortly before anthesis. Unusually Globba has zygomorphic style anatomy with only two abaxial vascular bundles, in contrast to most other Zingiberaceae, which possess three stylar traces. The ovary is unilocular and lacks septa. Bulbils have enclosing bracts and replace flowers in the lower part of the inflorescence; they consist of a shoot with an enlarged corky storage root forming the bulk of the propagule.     

Tree and grass rooting patterns in an African humid savanna

Abstract. Spatial and temporal soil partitioning between roots of the two savanna plant components, i.e. trees and grasses, were investigated in a West African humid savanna. Vertical root phytomass distribution was described for grass roots, large (> 2 mm) and fine (< 2 mm) tree roots, in open sites and beneath tree canopies. These profiles were established monthly over one year of vegetation growth. Natural ¹³C abundance measurement was used to determine the woody/herbaceous phytomass ratio in root samples. Tree and grass root distributions widely overlapped and both were mostly located in the top 20 cm of the soil. Grass root phytomass decreased with depth whereas woody root phytomass peaked at about 10 cm depth. No time partitioning was detected. These structural results do not support the hypothesis of soil resource partitioning between trees and grasses and are thus consistent with functional results previously reported.     

Polychlorinated biphenyls in alfalfa: Accumulation,sorption and speciation in different plant parts

The accumulation, chemical speciation and distribution of polychlorinated biphenyls (PCBs) were investigated in various parts of alfalfa. Moreover, the adsorption characteristics for PCB 28 by alfalfa and the influencing factors of the adsorption characteristics were studied. There were different degrees of PCB accumulation in alfalfa roots, root nodules and shoots. The decreasing order of the accumulation of PCBs in plant tissues was root nodules > roots > shoots, and the decreasing order of the total PCB contents was roots > shoots > root nodules, indicating that the roots were the main sink for PCB accumulation. There were three modes of PCB speciation in alfalfa roots and root nodules, comprising strong sorption (78%) and weak sorption (19%) on tissue surfaces and absorption within tissues (2%). The adsorption isotherms of PCB 28 indicate that the adsorption capacities of root nodules and shoots were both significantly higher than that of the roots. Both lipids and carbohydrates, and especially lipids, affected the PCB adsorption capacities of the tissues. These results may help in the elucidation of the mechanisms of sorption and accumulation of PCBs in the plants and their main influencing factors and thus contribute to the development of phytoremediation technologies for PCB-contaminated soils.     

Gas and Liquids in Intercellular Spaces of Maize Roots

Oils are spontaneously absorbed by gas-filled intercellularspaces (IS) in maize root cortex. The network of these spacesin living root sections was imaged by confocal laser scanningmicroscopy using a fluorescent solution of Nile red in oil.The gas volume fraction (GVF) of root segments was quantifiedby the increase in weight (differentiated zones) or tissue density(2–3 mm root tips) due to complete vacuum infiltration.Cooling to 6 °C or inhibition of oxidative phosphorylationdiminished the GVF of root tips but did not significantly affectthe GVF of differentiated root zones. The threshold pressuredifference for measurable infiltration of isolated root segmentsis lower (10 to 15  kPa) than the threshold for infiltrationof comparable zones of attached roots or of detached roots withthe cut surface sealed (>60 kPa). In the absence of an opencut, pressure-driven infiltration of the root cortex is acceleratedby microscopic fissures within the epidermal/hypodermal barrier.The GVF of the root cortex was reduced after transferring rootsfrom sugar solutions (0.1 to 0.3M ) to water. This points toefficient water transport from the medium to sugar-containingcortical cell walls through epidermal and hypodermal protoplasts.When 2-cm-long primary roots were vacuum infiltrated in situand then allowed to grow on aerated mineral medium for a further5 d, cortical IS of the originally infiltrated root bases remainedfilled with liquid but the subsequently grown apical root zoneshad a normal GVF. Copyright 1999 Annals of Botany Company Apoplastic and protoplasmic route, maize, infiltration, intercellular spaces, oil absorption, confocal laser scanning microscope, water transport, Zea mays L.     

Effects of liming on rhizosphere chemistry and growth of fine roots and of shoots of sessile oak (Quercus petraea)

Soil acidification can be detrimental to root growth and nutrient uptake, and liming may alleviate such acidification. In the following study, seedlings of sessile oak (Quercus petraea Liebl. M.) were grown in rhizotrons and subjected to liming (L) or gypsum (G) treatments and compared with the control (C). In order to study and interpret the impact of these calcium rich treatments on fine root development and tree growth, the following parameters were assessed: fine root biomass, fine root length, seedling development (height, diameter, leaves), seedling biomass, nutrient content of roots and seedlings, bulk soil and soil solution chemistry and rhizosphere soil chemistry. The results show that liming increased bulk soil pH, exchangeable Mg, Ca and the Ca/Al molar ratio, and decreased exchangeable Al, mainly in the A-horizon. Gypsum had a similar but smaller impact on exchangeable Al, Ca, H⁺ and the Ca/Al molar ratio in the A-horizon, but reacted with depth, so that exchangeable Mn, Mg and Ca were increased in the B-horizon. In the rhizosphere, the general pattern was determined by the treatment effects of the bulk soil. Most elements were more concentrated in the rhizosphere than in bulk soil, except for Ca which was less concentrated after liming or gypsum application. In the B-horizon rhizosphere pH was increased by the treatments (L > G,C) close to the root tips. Furthermore, the length of the zone with a positive root-induced pH increase was greater for the limed roots as compared with both the other treatments. Fine root growth was stimulated by liming (L > G,C) both in terms of biomass and length, whereas specific root length was not obviously affected apart from the indication of some stimulation after liming at the beginning. The live:dead ratio of fine roots was significantly higher in the limed rhizotrons as compared to the control (G not assessed), indicating lower mortality (higher longevity). Shoot growth showed greater lime-induced stimulation (L > G,C) as compared to root growth. As a result the shoot:root ratio was higher in the limed rhizotrons than in the control (L > G,C). Liming induced a higher allocation of P, S, Mg, Ca and K to the leaves, stem and twigs. Gypsum showed similar effects, but was only significant for S. Liming increased the foliar Ca/Al ratio by both increasing foliar Ca and decreasing foliar Al, whereas gypsum did not clearly improve foliar nutrition. This study suggests that a moderate application of lime can be successful in stimulating seedling growth, but that gypsum had no effect on seedling growth. It can be concluded that this lime-induced growth stimulation is directly related to the improved soil fertility status, and the alleviation of Al toxicity and acid stress, resulting in better foliar nutrition. The impact of liming on fine roots, as a consequence, was not limited to a stimulation of the total amount of fine roots, but also improved the root uptake performance. This revised version was published online in June 2006 with corrections to the Cover Date.     

Bryophyte County Floras

Abstract

Experimental study of Pohlia populations that regularly produce axillary bulbils in Britain has shown that there are seven species present. The distinguishing bulbil characters of field populations of each species are retained when the plants are cultivated under uniform conditions, and can be used as a valid taxonomic criterion. Variation in bulbil morphology within and between populations of P. proligera is due to a combination of genetic, environmental and developmental variation. Bulbils show considerable phenotypic plasticity in P. drummondii but in the other five species studied variation is largely or entirely developmen tal.     

Morphological and anatomical relationships of loblolly pine fine roots

 Suberized or brown roots have been traditionally considered secondary or woody tissues. The validity of using morphological features such as color to infer root anatomy for southern pines is questionable and unproven. The objectives of this study were (i) to establish relationships between root color, diameter, and developmental stage (i.e., primary or secondary tissues) for loblolly pine, (ii) to determine the percentages of primary and secondary brown roots by diameter class, and (iii) to use these percentages to make first order estimates of the amount of brown root length and surface area that is in the primary and secondary developmental stages for sampled roots of a semi-mature loblolly pine stand. ”Unsectioned” roots were collected by coring to a 25-cm depth 3 times a year and measuring roots for length and surface area by diameter class. ”Sectioned” roots were sampled from a one-time core and from periodic grab samples. These roots were sectioned and characterized by their color, diameter and developmental stage. Diameters of sectioned roots (n=353) ranged from 0.21 to 8.24 mm. White and orange roots ranged from 0.23 to 2.50 mm, while brown roots spanned the range. White roots were developmentally primary, whereas orange/brown roots were either primary (from 0.21 to 2.50 mm), secondary (from 0.33 to 8.24 mm), or in transition (from 0.27 to 0.76). Total live root length of the sampled stands was estimated to be composed of 38% primary tissue, 58% secondary tissue, and 4% transition tissue. Lastly, neither root color nor diameter was a reliable predictor of developmental stage unless roots were white (primary), or orange/brown and >2.5 mm in diameter (secondary). Received: 30 June 1997 / Accepted: 28 January 1998     

Development of the root crown in some conifers

Summary Large and small stumps of 3 spruce species, a pine, a fir and hemlock were pulled up from 15 year plantations. Roots originating from the root crown were counted (<0.5 cm) or, if larger, their diameters were measured, taper, eccentricity, compass bearing, inclination, position on the rootstock and basal age (by annual rings). The sequence of events leading to the developed skeletal root system was reconstructed. Hypotheses to account for the distinct populations of roots were examined. In all species, major roots originated between 2 and 7 yrs after planting and grew thicker and more eccentric at their bases than minor roots of the same age. Positively geotropic roots were distinct from plagiogeotropic roots.     

Gravitropism of Primary Roots of Zea mays L. at Different Displacement Angles

Primary roots of Zea mays were oriented at various angles fromthe vertical ranging from 99° to 1° and their subsequentbending analysed from filmed records. The maximum rate of bendingand the time before bending commenced both varied two-fold,but showed no correlation with the initial angle of tip displacement.Roots orientated to small initial angles (< 40°) oftenovershot the vertical and proceeded to oscillate around thisorientation, whereas roots oriented to large initial angles(> 60°) often failed to achieve the vertical. Roots inthis latter group resumed bending after an indeterminate time,or did so immediately after a second displacement of their tip,showing that they were not intrinsically unable to bend. Theapparently spontaneous resumption of bending after a temporaryplagiogravitropic phase is suggested as being due to noise inthe graviperception system in the root cap. The tips of rootsgrowing vertically downwards showed oscillatory bending movementsup to 10° either side of vertical. This angle correspondsto the minimum angle of displacement which induces gravitropicbending. Only when roots were oriented 10-20° from verticaldid they begin unequivocally to show a gravitropism since atsuch angles the deflection of their tips exceeded that due totheir natural oscillation.Copyright 1993, 1999 Academic Press Gravitropism, roots, Zea mays