Factors Affecting the Growth of Daughter Bulbs in the Tulip

The relative growth rate (R_w) of daughter bulbs of the tulipcultivar Rose Copland was remarkably constant during the springperiod of growth in four seasons at two sites. The Q₁₀ of R_wwas 2.2, and the long period of constant R_w is attributed tocompensation of a fall with age by increasing temperatures inthe spring. Final bulb weight differed among four cultivarsbecause of differences in R_w and in initial daughter bulb weights. Partial defoliation reduced R_w roughly in proportion to theleaf area removed, and removal of mother bulb scales resultedin reduced leaf area, fewer daughter bulbs, and a lower daughterbulb R_w. Heat-treatment of mother bulbs before planting (blindstoken)killed the flower within the bulb, inactivated the apical dominanceexerted by the flower, and resulted in a higher initial daughter-bulbweight at the start of the spring period of exponential growth.The R_w of heat-treated and control daughter bulbs were not different,neither were the leaf areas, so it is assumed that final daughter-bulbweights were higher following treatment because of increasedsink strength. The economic implications of these findings arediscussed.     

The Initiation and Growth of Narcissus Bulbs

Plants of two Narcissus cultivars were dissected periodicallyover a year to study initiation, growth, and dry-weight changesof flowers and daughter bulb units in relation to the positionof the daughter bulbs in the branching system. A Narcissus bulbis a branching system comprising terminal and lateral bulb units;the former bear both terminal and lateral bulb units, but lateralscontain only terminal ones. This basic pattern may be modifiedby the failure of lateral bulb units or the development of supernumeraryones. Differences in bulb unit size, in scale and leaf number,and in flowering, related to position in the branching system,are probably due to the initiation of lateral bulb units somemonths after terminal ones. This affects the growth and behaviourof the laterals and daughter bulb units borne by these lateralsin the next generation. Bulb units live about four years; after the loss of their leafblades and flowers, the scales and leaf bases act as storageorgans whose weight increases or decreases according to thecarbohydrate status of the plant. Growth of bulb units showsan externally controlled alternation of rapid and slow growth.This is likely to be an effect of day length on bulbing.     

Photoperiod and Temperature Affect Sprouting of Onion bulbs (Allium cepa L.)

The effects of photoperiod and temperature on sprouting wereinvestigated using three cultivars of onion which exhibit differentresponses to daylength during bulb growth. Two photoperiods(8 h and 14 h) and two constant temperatures (15 C and 20 C)were used. Two cultivars showed enhanced sprouting and leafemergence in short days, and one sprouted poorly in all conditions.In long days, some plants showed growth of new bulbs withinthe old one, without sprouting. Photomorphogenesis, photoperiodism, Allium cepa L, onion, dormancy, bulb sprouting     

The Role of Growth Substances in the Regulation of Onion Bulb Dormancy

The correlation between sprouting and changes in endogenousgrowth substances was investigated in stored onion bulbs (Alliumcepa c.v. Rijnsburger and Lancastrian). In the main experimentsbulbs were removed from store at approximately fortnightly intervals,samples were assessed for percentage sprouting and non-sproutingbulbs were either extracted for hormone assay or treated withgrowth substances in an attempt to induce sprouting. In otherexperiments the hormone content of bulbs at different stagesof sprouting was assessed. Growth-inhibitor and gibberellinactivity decreased before sprouting, but there was an increasein gibberellin and auxin activity as sprouting commenced. Gibberellinactivity was highest in bulbs with well-developed sprouts whereasauxin activity occurred mainly in bulbs in which early sproutdevelopment was visible only on their being cut open. Therewas no conclusive evidence that bulb dormancy could be brokenby application of the gibberellins GA₃ and GA_4/7, or the auxin1-naphthylacetic acid (NAA). Maleic hydrazide (MH) completelyinhibited root and sprout development but the growth retardant(2-chloroethyl)trimethylammonium chloride (CCC) was mainly effectivein reducing root development and sprouting was only slightlyinhibited.     

Biomass Allocation of Erythronium americanum Populations in Different Irradiance Levels

The forest herb Erythronium americanum was collected from threeNew Jersey habitats and grown in 100%, 43%, or 23% glasshouselight to determine if biomass allocation responses to irradiancediffered between populations. Shaded plants had longer petioles,produced fewer runners, and had greater specific leaf areasthan unshaded plants; these responses did not differ significantlyamong populations. All populations produced smaller bulbs whengrown in shade. Over all light treatments, mean leaf area ofplants from a stable, established site was greater than thatof plants from others. In 100 % light, these plants had thegreater bulb growth. The simple architecture of Erythroniummay constrain morphological responses to the light environment. Erythronium americanum(Ker.), forest herb, biomass allocation, light intensity     

Changes in Endogenous Gibberellin Levels in Tulipa Bulblets during Ontogeny

The endogenous gibberellin activity of Tulipa gesneriana cv.Apeldoorn bulbiets of field-grown mother bulbs was determinedduring ontogeny by paper and gas-liquid chromatography and thedwarf pea bioassay. It was shown that the gibberellin activityof bulblets increased dramatically in November-March and declinedsharply in April-July. The increase in gibberellin-like substanceswas considered to be derived primarily by synthesis within thebulblets with possible contributions via translocation fromthe mother bulb scales, roots and shoots. Gibberellin A₁₃ wastentatively identified by gas-liquid chromatography but theother components of the bulbiet extracts remained unidentified.     

Neoseiulus paspalivorus,a predator from coconut,as a candidate for controlling dry bulb mites infesting stored tulip bulbs

The dry bulb mite, Aceria tulipae, is the most important pest of stored tulip bulbs in The Netherlands. This tiny, eriophyoid mite hides in the narrow space between scales in the interior of the bulb. To achieve biological control of this hidden pest, candidate predators small enough to move in between the bulb scales are required. Earlier experiments have shown this potential for the phytoseiid mite, Neoseiulus cucumeris, but only after the bulbs were exposed to ethylene, a plant hormone that causes a slight increase in the distance between tulip bulb scales, just sufficient to allow this predator to reach the interior part of the bulb. Applying ethylene, however, is not an option in practice because it causes malformation of tulip flowers. In fact, to prevent this cosmetic damage, bulb growers ventilate rooms where tulip bulbs are stored, thereby removing ethylene produced by the bulbs (e.g. in response to mite or fungus infestation). Recently, studies on the role of predatory mites in controlling another eriophyoid mite on coconuts led to the discovery of an exceptionally small phytoseiid mite, Neoseiulus paspalivorus. This predator is able to move under the perianth of coconuts where coconut mites feed on meristematic tissue of the fruit. This discovery prompted us to test N. paspalivorus for its ability to control A. tulipae on tulip bulbs under storage conditions (ventilated rooms with bulbs in open boxes; 23 °C; storage period June–October). Using destructive sampling we monitored predator and prey populations in two series of replicated experiments, one at a high initial level of dry bulb mite infestation, late in the storage period, and another at a low initial dry bulb mite infestation, halfway the storage period. The first and the second series involved treatment with N. paspalivorus and a control experiment, but the second series had an additional treatment in which the predator N. cucumeris was released. Taking the two series of experiments together we found that N. paspalivorus controlled the populations of dry bulb mites both on the outer scale of the bulbs as well as in the interior part of the bulbs, whereas N. cucumeris significantly reduced the population of dry bulb mites on the outer scale, but not in the interior part of the bulb. Moreover, N. paspalivorus was found predominantly inside the bulb, whereas N. cucumeris was only found on the outer scale, thereby confirming our hypothesis that the small size of N. paspalivorus facilitates access to the interior of the bulbs. We argue that N. paspalivorus is a promising candidate for the biological control of dry bulb mites on tulip bulbs under storage conditions in the Netherlands.     

Changes in the Endogenous Growth Regulators in Bulbous Iris in Bulb-forming and Nonbulb-forming Aspects

Bulblets of 3–4 g of Dutch iris (Iris hollandica) cv.Dominator stored at 20°C and then grown at 15°C developedonly three or four leaves and bulbs formed at the base of eachleaf, whereas when grown at 25°C, they continued to growvegetatively with the development of 10 true leaves and didnot form any new bulb. This demonstrated that bulb formationin bulbous plants can be controlled by environmental factors.The levels of both abscisic acid and auxin activities increasedduring growth under the bulb-forming condition whereas onlyauxin activity increased under the nonbulb-forming condition.The coexistence of both abscisic acid and auxin seems to beessential in the processes of bulb formation. (Received August 22, 1980; Accepted November 26, 1980)     

Totipotency in Tissue Explants and Callus of Some Members of the Liliaceae, Iridaceae, and Amaryllidaceae

In vitro responses of twelve species of bulbs and conns werecompared. Plantlets could be induced directly without interveningcallus on stem tissue in nine species, on ovary tissue in fivespecies, and on leaf tissue in four species. In Gladiolus, Hyacinthus,Muscari, Ornithogahim, and Scilla plantlets were formed withoutgrowth factors added to the Murashige and Skoog medium. In Hippeastrum,Schizoslylis, Sparaxis, and Ipheion auxin was required. No plantletscould be induced directly on explants of growing tissue of Freesia,Tulipa, or Narcissus. Adventitious plantlets could be inducedon pieces of bulb or corm from ten species but such materialwas difficult to free from contamination. Callus was obtained from all species except Tulipa and Hippeastrum.Plantlets could be regenerated from callus except that of Gladiolus,Sparaxis, and Schizostylis. Differences in responses of the twelve species tend to cut acrossthe three families and no simple relation is evident betweenthe natural rate of vegetative increase and the in vitro behaviour.     

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.     

Aspects of the Comparative Physiology of Ranunculus bulbosus L. and Ranunculus repens L.: I. Response to Nitrogen

Ranunculus bulbosus and R. repens are closely related taxonomicallyand coexist in the same general habitat but differ in that R.bulbosus only exceptionally shows vegetative multiplicationcontrasting with R. repens which reproduces extensively by meansof long branched stolons which root and form new daughter plantsat the nodes. The response of these two species to nitrogen was similar inthat an increase from 10 to 50 ppm nitrogen gave rise to greatlyincreased total dry weight and higher shoot to root ratios,whereas further nitrogen increase from 50 to 200 ppm had littleeffect. However, they differed in that at low nitrogen levelsthe perennating organ (corm) took priority in R. bulbosus withsexual reproduction assuming greater importance at the highernitrogen levels, whereas in R. repens the main storage organ(the stock) was not especially favoured at low nitrogen levelsreflecting perhaps the differences between this structure andthe corm in R. bulbosus. In R. repens the greatest responseto nitrogen was shown by the stolons although this was due toincreased branching and a higher dry weight per unit lengthat the higher levels of nitrogen (50 and 200 ppm). Primary stolonlength was not greatly affected by nitrogen level. Dry weightsof flowers and fruits in R. repens were very much lower thanin R. bulbosus. It is suggested that in R. bulbosus the emphasis lies firstin replacement of the parent at approximately the same siteby a daughter from the new corm and secondly in sexual reproductivespread. In R. repens lateral spread by vegetative propagationtakes precedence over all else and under low nitrogen conditionsbranching is sacrificed and energy is diverted to maintaininglong primary stolons. These, in turn, have the potential forproducing daughter plants at some distance from the parent andthis may be of considerable importance in field situations sincethe daughter may come to occupy more favourable sites than theparent.     

Catecholaminergic contributions to the neuronal machinery of the olfactory bulb: aftoradiographic, immunohistochemical and evoked feild potential studies

aftographic exeperiments on the localization of radiolabelednoradrenaline, dopamine and dopa, as well as immunohistochemicalstudies on hydroxylase-like activity, are summarized and comparedin both rat and turtle olfactory bulbs. Evoked field potentialstudies on effects of dopamine are also discussed. The histochemicalstudies suggest that dopaminergic periglomerular neurons arethe most significant cellular component of the catecholaminergicsystem in the olfactory bulb of both species. Scattered fluorescentcell group was also present in the internal plexiform layerand superficial granule cell layer of the turtle olfactory bulb.Other fibres, not related to intrinsic bulbar neuronal cellbodies, were also labeled, mostly in the granule cell layerbut also in the external plexiform layer. These might belongto a centrifugal catecholaminergic system from brain stem neurons.In the in vitro turtle olfactory bulb, dopamine and apomorphinedepressed the amplitude of field potentials evoked by a singlevolley in the olfactory nerve or lateral olfactory tract, andreduced the depression and latency of reponses when paired volleywere delivered. It is suggested that catecholaminergic systemsplay a key role in modulating mitral cell activity through actionsin both superficial (glomerular) and deep (granule) layers.This may involve direct actions, or other, non-catecholaminergicinterneurons.     

Phosphate Uptake and Transport by Roots and Stolons of Intact White Clover (Trifolium repens L.)

The extent to which phosphate can be absorbed directly fromthe outer medium by stolon internodes and contribute to thetotal accumulation of phosphate by intact plants of white clover(Trifolium repens L. cv. Blanca) was assessed in hydroponicexperiments in a controlled environment room. The uptake ofphosphate by intact roots or stolons was measured by sealinga segment (6-0 mm long) across a flow-cell in which ³²P-labellednutrient solution was circulated for 24 h, the rest of the rootsystem receiving unlabelled nutrient solution. The rate of uptakeof phosphate (µmol g^–1 d^–1 dry wt. basis)by roots was more than 300 times that by intact stolons. Pretreatmentof stolons by gentle abrasion to remove cuticle, so as to simulatethe condition of stolons in the field, increased the uptakeof phosphate 7-fold compared with that of intact stolons. However,the potential of stolons to contribute to the P status of whitedover in the field was calculated to be small (5%). When an incision was made through the hypodermal layer of stolons,the rate of phosphate uptake greatly increased, attaining 71%of that by root segments. This increase, which was greater athigher phosphate concentrations, indicates that the suberi.zedhypodermis constitutes a major barrier to the influx of phosphatein the stolon. After withholding phosphate for different time intervals, thesubsequent rate of phosphate uptake by roots was increased 2-3-foldafter 2 d phosphate deprivation and 3-4-fold after 6 d or 13d phosphate deprivation. A higher proportion of absorbed phosphatewas transported to shoots in phosphate-deprived plants. After1 d of uptake following restoration of the phosphate supply,the concentrations of labelled phosphate in shoots were greaterthan in control plants, although the concentrations of labelin roots was less. However, the rate of uptake of phosphateby stolons, following deprivation, was not significantly increased.These results suggest that the mechanism regulating the enhancedrate of phosphate loading into the xylem, initiated by a periodof phosphate deprivation, is specific to roots and is not inducedin stolons. The results are discussed in relation to the growth and acquisitionof phosphate by white clover in the field. Key words: Nutrient deficiency, phosphate, stolons, transport (ions), Trifolium repens     

Basal Plate Tissue in Narcissus Bulbs and in Shoot Clump Cultures: Its Structure and Role in Organogenic Potential of Single Leaf Cultures

Light microscopy demonstrated that the apparently amorphous,achlorophyllous tissue at the base of in vitro shoot clump cultureof Narcissus was comparable in structure to the basal plateof Narcissus bulbs. Both had very complex vascularisation andsmall, densely packed parenchymatous cells. In shoot clump cultures, primordia were produced by meristematiczones at the surface of this achlorophyllous tissue, very closeto the base of leaves. Single leaf units excised from the invitro shoot clump cultures with a wedge of basal achlorophylloustissue were highly organogenic when used as secondary explantsfor in vitro culture of Narcissus. No organogenesis occurredin the absence of the leaf base and achlorophyllous (basal plate)tissue and little organogenesis occurred unless the leaf baseand basal plate tissue were immersed in the culture medium (i.e.explants inoculated into liquid medium or upright in agar-solidifiedmedium). After two 5-week culture passages in liquid medium, more thanfive leaves were produced per leaf base inoculated. Thus rapidmicropropagation of Narcissus can be achieved using only thebase of single leaf units excised from shoot clump cultures.Copyright1993, 1999 Academic Press Anatomy, basal plate, bulb, in vitro, leaf culture, Narcissus, organogenesis     

Bulbil production in Narcissus: the effect of temperature and duration of storage on bulb unit development and subsequent propagation by twin-scaling

The growth of bulb components and carbohydrate content of Narcissus bulbs was followed during storage at 5–30 ^oC, and the productivity of twin-scale propagules cut from bulbs after these storage treatments was studied. Growth of daughter bulb unit components was greatest during storage at 9–15 ^oC, and greatly inhibited at 30 ^oC. Concentrations of soluble sugars were highest following 30 ^oC storage, and higher in intermediate bulb scales than in outer or inner scales. Bulbil production on twin-scales was greatest on propagules cut from the intermediate scales, and in bulbs previously stored at 15 or 30 ^oC.     

Contribution from Stolons and Roots to Estimates of the Total Amount of N2Fixed by White Clover (Trifolium repensL.)

Growth and N-accumulation rates in leaves, stolons and rootsof individual white clover plants were studied in three experimentsusing two methods. In a growth chamber experiment, the relativedifferences between tissues were found to be almost constantfor a wide range of clover plant sizes. The stolon dry matter(DM) production was 56% and the root DM production 40% of theDM production in leaves. The N yield of stolons was 30% whileN yield in roots was 34% of N yield in leaves. The effect ofN application on these relations was investigated in a glasshouseexperiment. Application of N reduced the root:shoot N ratiofrom 0.50 to 0.28, whereas the stolon+root:leaf N ratio (i.e.for abovevs.below cutting-height tissues) was only reduced from0.97 to 0.80. In a field trial with two contrasting N regimes,growth and N accumulation were measured on individual cloverplants. Dinitrogen fixation was estimated by¹⁵N isotope dilutionbased on analysis of leaves-only or by including stolons. Usingleaves-only did not affect the calculation of percentage ofclover N derived from N₂fixation (% Ndfa) since the¹⁵N enrichmentwas found to be uniform in all parts of the clover. A correctionfactor of 1.7 to account for N in below cutting-height tissueis suggested when N₂fixation in white clover is estimated byharvesting the leaves only.Copyright 1997 Annals of Botany Company Leaves; N accumulation; N₂fixation; ¹⁵N isotope dilution; pastures; roots; root/shoot ratio; stolons; Trifolium repensL.; white clover     

Water Relations of White Clover (Trifolium repens): Water Potential Gradients and Plant Morphology

Water potential, osmotic potential, pressure potential and relativewater content were measured in stolons and leaves of white cloverplants grown under a range of conditions of water supply andevaporative demand. The importance of adventitious roots fromthe nodes was examined. Gradients along stolons were alwaysextremely small, of the order of only 01 MPa. Stolon up waterpotential was representative of plant water status regardlessof stolon length, presence/absence of nodal roots, degree ofwater stress and evaporative demand. It is concluded that waterconduction along stolons was very good. Gradients were foundto exist along petioles; they may have a greater resistanceto water flow than stolons. The relationship between water fluxand stem anatomy, and the importance of differential flow ratesthrough stolons and petioles to plant behaviour during waterstress, are discussed. Trifolium repens L., white clover, water relations     

Respiration Rate and Mitochondrial Activity in Iris Bulbs

The oxygen uptake of iris bulbs (Iris×hollandica‘Wedgwood’) which had been stored dry at 30 C (“retarded’ bulbs) was low (10 μmol O₂ per h and bulb), the oxygen uptake of the intact bulb, the three outer fleshy scales and the remaining central part of the bulb increased three- to fourfold, nearly twofold and fourfold, respectively. Mitochondria were isolated from the scales of retarded and activated bulbs and their oxygen consumption with succinate, l -malate (plus pyruvate). x-ketoglutarate and NADH as substrate was measured polarographically. The oxidative capacity of mitochondria isolated from the scales of activated bulbs was only slightly higher than that from retarded bulbs when calculated on a tissue basis. No difference was found between the phosphorylation efficiency, respiratory control, cytochrome c deficiency, succinate dehydrogenase, malate dehydrogenase, succinate-cytochrome c rductase, NADH-cytochrome c reductase and cytochrome oxidase activity of the retarded and activated bulbs. The increase in the in vitro oxygen uptake of the scales after transition from 30 to 13 C was not accompanied by an equal increase in the oxidative capacity of their mitochondria suggesting that they are not responsible for this rise in oxygen uptake.     

Tuberization in Potato at High Temperatures: Gibberellin Content and Transport from Buds

Warm temperatures (35°C day/30°C night) which inhibittuberization in potato (Solanum tuberosum L., cv. Sebago) increasedgibberellin activity in crude extracts from buds, but not frommature leaves, as determined by the lettuce hypocotyl bioassay.Changes in the growth of tubers and stolons indicate the occurrenceof basipetal movement of GA₃ applied to the terminal bud ora mature leaf. ¹⁴C labelling from GA₃ or mevalonic acid injectedjust below the terminal bud was recovered in the lower shoot,stolons and tubers, but the amount transported was greater atcool temperatures (20/15°C). It is concluded that high temperaturespromote the synthesis of gibberellin in the buds rather thantransport to the stolons. Solanum tuberosum L., potato, tuberization, gibberellin     

Stomatal responses to a range of variables in two tropical tree species grown with CO2 enrichment

Seedlings of Maranthes corymbosa (Blume) and Eucalyptus tetrodonta(F. Muell) were grown with or without CO₂ enrichment (700µmolCO₂ mol^–1 The response of stomatal conductance (g₂) toleaf drying, exogenous absclslc acid and calcium ions was investigatedin M. corymbosa. Reciprocal transfer experiments were also conductedwhereby plants were grown in one treatment and then transferredto the other before g, was measured. Stomatal conductance in M. corymbosa was more sensitive (a greaterpercentage decline in g₂ per unit percentage decline in leaffresh weight) to leaf water status under conditions of CO₂ enrichmentcompared to ambient conditions. However, the rate of reductionof g₂ in response to exogenous abscisic acid was not influencedby CO₂ treatment. In contrast, the rate of reduction of g₂,in response to exogenous CaCl₂ was decreased under conditionsof CO₂ enrichment. Reciprocal transfer experiments showed that expo sure to CO₂enrichment results in a short-term, reversible decline in g₂,as a result of decreased stomatal aperture and a long-term,irreversible decline in g₂ as a result of a decreased stomataldensity. Seedlings of E. tetrodonta were used to investigate the responseof g₂ to light flux density, leaf-to-air vapour pressure difference(LAVPD), leaf internal CO₂ concentration (C₁ and temperature.Reciprocal transfer experiments were also conducted. CO₂ enrichmentdid not influence the pattern or sensitivity of response ofg to LAVPD and C in E. tetrodonta. In contrast, the slope ofthe response of g₂, to temperature decreased for trees grownunder elevated [CO₂]_a conditions and the equilibrium g₂ attainedat saturating light was also decreased for plants grown underelevated [CO_2a. conditions. Key words: Stomata, elevated CO₂, tropical trees