Bud Structure in Relation to Shoot Morphology and Position on the Vegetative Annual Shoots of Juglans regia L. (Juglandaceae)

The length and basal diameter of all lateral and terminal budsof vegetative annual shoots of 7-year-oldJuglans regia treeswere measured. All buds were dissected and numbers of cataphylls,embryonic leaves and leaf primordia were recorded. Each axillarybud was ranked according to the position of its associated leaffrom the apex to the base of its parent shoot. Bud size andcontent were analysed in relation to bud position and were comparedwith the size and number of leaves of shoots in equivalent positionswhich extended during the following growing season. Length andbasal diameter of axillary buds varied according to their positionon the parent shoot. Terminal buds contained more embryonicleaves than any axillary bud. The number of leaves was smallerfor apical and basal axillary buds than for buds in intermediatepositions on the parent shoot only. All new extended shootswere entirely preformed in the buds that gave rise to them.Lateral shoots were formed in the median part of the parentshoot. These lateral shoots derived from buds which were largerthan both apical and basal ones. Copyright 2001 Annals of BotanyCompany Juglans regia L., Persian walnut tree, branching pattern, preformation, bud content, shoot morphology     

Acorn production is linked to secondary growth but not to declining carbohydrate concentrations in current-year shoots of two oak species

In trees, reproduction constitutes an important resource investment which may compete with growth for resources. However, detailed analyses on how growth and fruit production interact at the shoot level are scarce. Primary canopy growth depends on the development of current-year shoots and their secondary growth might also influence the number and size of fruits supported by them. We hypothesise that an enhanced thickening of current-year shoots is linked positively to acorn production in oaks. We analysed the effect of acorn production on shoot growth of two co-occurring Mediterranean oak species with contrasting leaf habit (Quercus ilex, Quercus faginea). Length and cross-sectional area of current-year shoots, apical bud mass, number of leaves and acorns, xylem and conductive area, number of vessels of acorn-bearing and non-bearing shoots were measured in summer and autumn. Nitrogen and carbohydrates analyses were also performed in stems and leaves of both shoot types. Stem cross-sectional area increased in acorn-bearing shoots when compared with non-bearing shoots for both species and such surplus secondary growth was observed since summer. In bearing shoots, the total transversal area occupied by vessels decreased significantly from basal to apical positions along the stem as did the xylem area and the number of vessels. Leaves of bearing shoots showed lower nitrogen concentration than those of non-bearing shoots. Carbohydrate concentrations did not differ in stems and leaves as a function of the presence of acorns. Such results suggest that carbohydrates may preferentially be allocated towards reproductive shoots, possibly through enhanced secondary growth, satisfying all their carbon demands for growth and reproduction. Our findings indicate that acorn production in the two studied oaks depends on shoot secondary growth.     

Allocation of resources within mountain birch canopy after simulated winter browsing

As a response to browsing, birches are known to produce fewer but larger, more nutritious leaves, with enhanced palatability for herbivores. We simulated winter browsing in ramets of mountain birch ( Betula pubescens ssp. czerepanovii ) to find out whether it decreases subsequent foliage biomass and alters the number and type of shoots. After removal of a considerable proportion of buds (up to 35%) in late winter, the birches were able to compensate for the lost leaf biomass in the following summer; there were no differences in total leaf biomass between winter-clipped and control ramets. This indicates that foliage growth was limited by the total amount of stored resources, not by the number of buds. Depending on the position of the buds removed, different mechanisms were responsible for the compensation. After removal of apical buds, the number of leaves decreased significantly but leaves were larger than in control ramets. Removal of the same mass of basal buds – containing similar amount of carbohydrates and proteins as in the treatment removing apical buds – activated dormant buds, especially in apical locations, so that leaf number was similar as in the controls; consequently, size of individual leaves increased only slightly. Thus, while the total leaf biomass in a tree seems to be limited by resources from source organs, the distribution of resources among different canopy sections is controlled by their relative sink strengths. In terms of leaf biomass, apical parts are able to compensate for bud loss by increasing shoot number, basal parts only by increasing leaf size.     

Role of phytohormones in organogenic ability of elm multiplicated shoots

The study presents the comparative analyses of endogenous contents of auxin (IAA), cytokinins (CKs), polyamines (PAs), and phenolic acids (PhAs) in apical and basal parts of elm multiplicated shoots with regard to the organogenic potential. The shoot-forming capacity was higher in the apical part than in the basal part. However, the timing of root formation was in the apical type of explant significantly delayed (compared with the organogenic potential of basal part). Significantly higher contents of free bases, ribosides and ribotides of isopentenyl adenine, zeatin and dihydrozeatin that were found in the apical segments, might be considered as the most important factor affecting in vitro shoot formation. The content of endogenous free IAA was approximately three times higher in the basal shoot parts than in the apical parts. The amounts of putrescine and spermidine were higher in the apical part which generally contains less differentiated tissues than the basal part of shoot. The predominant PhA in both types of explants was caffeic acid, and concentrations of other PhAs decreased in the following order: p-coumaric, ferulic, sinapic, vanillic, chlorogenic, p-hydroxybenzoic and gallic acids. The contents of all determined PhAs in their free forms and higher contents of glycoside-bound p-coumaric, ferulic and sinapic acids, precursors for lignin biosynthesis, were found in the basal parts.     

Testing a new hypothesis: plant vigor and phylloxera distribution on wild grape in Arizona

Summary Longer, meaning more vigorous, shoots of a wild grape clone (Vitis arizonica) were more susceptible to attack by second and third generations of leaf-galling grape phylloxera,Daktulopsphaira vitifoliae, as the growing season progressed. Although there was no significant difference in mean shoot length between attacked and unattacked shoots within a clone at the beginning of shoot elongation, attacked shoots were significantly longer than unattacked shoots when elongation had ceased (P<0.01). Also, long attacked shoots had a significantly greater population of phylloxera galls than short attacked shoots (P<0.01) as the season progressed. The phylloxera population on long shoots increased rapidly while the population on short shoots remained the same. Longer shoots also produced significantly more axillary shoots than shorter shoots as the season progressed (P<0.001), and the number of axillary shoots accounted for 66 percent of the variance in number of attacked leaves on a shoot. Experimental evidence showed that there was a significantly greater percentage of available leaves attacked on long shoots than on short shoots (P<0.05) and the leaves on long shoots generally had a greater number of galls per leaf. The relationship between shoot length and probability of attack was also tested by comparing shoots lengths of 10 attacked clones and 10 unattacked clones at a second location. Mean shoot lengths of attacked clones were significantly longer than mean shoot lengths of unattacked clones (P<0.05), and mean shoot lengths of attacked shoots within a clone were significantly longer than unattacked shoots (P<0.001). Longer shoot length accounted for 81 percent of the variance in probability of attack. The reason for this pattern of attack was that long shoots produced newly expanding leaves over a longer time during the growing season and multivoltine phylloxera require undifferentiated tissue to initiate gall formation. Patterns of attack within a shoot were characterized by an uneven distribution of galls among leaves. This was due to development time between generations and the current availability of undifferentiated tissue at times of colonization. This study supports the hypothesis that some herbivore species are favored more by vigorous plants than by stressed plants.     

In vitro regeneration of plantlets in Brassica alboglabra

Different vegetative parts of Brassica alboglabra seedlings and mature plants were used as explants in culture.A high frequency (60–100%) of shoot regeneration was obtained from hypocotyl explants, nodal stem segments, internodal segments and shoot apices cultured on Murashige-Skoog basal medium. Addition of 6-benzylaminopurine and kinetin increased the average number of shoots per explant. When detached and transferred to basal medium, the shoots readily developed roots. Regenerated plantlets could be successfully transplanted in soil.     

Plant regrowth response to a stem-boring insect: a swift moth-willow system

To examine plastic willow regrowth response to herbivory, we studied the effect of a boring insect, the swift moth Endoclita excrescens (Hepialidae: Lepidoptera), which does not remove apical meristems, on shoot growth in three willow species—Salix gilgiana, S. eriocarpa, and S. serissaefolia−by direct observations and experiments in the field. We hypothesized that the stem-boring could initiate new lateral bud activation, and result in secondary shoot regrowth without the removal of the primary apical meristems. There were significantly more lateral shoots on naturally attacked than unattacked stems, and a significant positive correlation between lateral shoot density and the number of swift moth tunnels per tree was observed for all three willow species. Artificial boring, and larval infestation, resulted in an increase in the number of lateral shoots, but did not affect growth of current-year shoots. The length of lateral shoots differed between species, being significantly longer in S. gilgiana than S. eriocarpa and S. serissaefolia. The results of this study show that compensatory regrowth can result even if herbivory does not remove the apical meristem. We argue that this type of plant compensatory response is probably widespread, given that the stem-boring is a common feeding type of insect herbivores.     

Patterns of leaf size and morphology in relation to shoot length in Tsuga canadensis

Summary Pattern of change in leaf character was assessed along the length and around the circumference of Tsuga canadensis (L.) Carr. shoots of different length classes. Leaf size decreased, and number of leaves per unit length of shoot increased, with decrease in shoot length. Acropetally along the shoots, lamina length and width decreased, relative petiole length increased, apices became more pointed, and leaf margins bore more teeth. Around the shoot, from upper to side, and to lower surfaces, leaf size and number of marginal teeth in the proximal halves of leaves increased. These patterns were related both to production of preformed versus neoformed leaves, though their separation was indistinct, and to secondary orientation of leaves by twisting in their petiolar regions into two major bi-lateral ranks. An additional minor rank occurred along the upper surfaces of the shoots where secondary orientation of the leaves was minimal. Surface features of leaves did not differ in any obvious manner. Leaves on sylleptic shoots, which by definition were all neoformed, exhibited similar patterns, but were generally smaller than those on their parent shoots.     

Quantitation of endogenous levels of IAA,IAAsp and IBA in micro-propagated shoots of hybrid chestnut pre-treated with IBA

Endogenous levels of indole-3-acetic acid (IAA), indole-3-acetylaspartic acid (IAAsp) and indole-3-butyric acid (IBA) were measured during the first 8 d of in vitro rooting of rootstock from the chestnut ‘M3’ hybrid by high performance liquid chromatography (HPLC). Rooting was induced either by dipping the basal ends of the shoots into a 4.92-mM IBA solution for 1 min or by sub-culturing the shoots on solid rooting medium supplemented with 14.8-μM IBA for 5 d. For root development, the induced shoots were transferred to auxin-free solid medium. Auxins were measured in the apical and basal parts of the shoots by means of HPLC. Endogenous levels of IAA and IAAsp were found to be greater in IBA-treated shoots than in control shoots. In extracts of the basal parts of the shoots, the concentration of free IAA showed a significant peak 2 d after either root inductive method and a subsequent gradual decrease for the remainder of the time course. The concentration of IAAsp peaked at day 6 in extracts of the basal parts of shoots induced with 14.8-μM IBA for 5 d, whereas shoots induced by dipping showed an initial increase until day 2 and then remained stable. In extracts from basal shoot portions induced by dipping, IBA concentration showed a transient peak at day 1 and a plateau between day 2 and 4, in contrast to the profile of shoots induced on auxin-containing medium, which showed a significant reduction between 4 and 6 d after transferred to auxin-free medium. All quantified auxins remained at a relatively low level, virtually constant, in extracts from apical shoot portions, as well as in extracts from control non-rooting shoots. In conclusion, the natural auxin IAA is the signal responsible for root induction, although it is driven by exogenous IBA independently of the adding conditions.     

THE PHYSIOLOGICAL BASIS OF MORPHOLOGICAL POLARITY IN REGENERATION. I

1. In Bryophyllum calycinum two apical leaves suppress the shoot formation in all the dormant buds situated basally from the leaf; one apical leaf suppresses the shoot formation in the basal buds situated in the same half of the stem where the leaf is, and, if one-half of the petiole of such a leaf is removed, the growth of basal buds in one quadrant of the stem is suppressed. 2. This inhibitory influence of a leaf upon shoot formation in the basal part of a stem is diminished or disappears when the mass of the leaf is reduced below a certain limit. 3. The inhibitory influence of an apical leaf upon the growth of shoots in horizontally suspended stems is greater when the leaf is on the upper than when it is on the lower side of the stem. 4. All these facts suggest the possibility that the inhibitory influence of the leaf upon shoot formation is due to inhibitory substances secreted by the leaf and carried by the sap from the leaf towards the base of the stem. 5. An apical leaf accelerates root formation in the basal part of a stem and this accelerating effect increases with the mass of the leaf. 6. This inhibitory influence of a leaf upon shoot formation and the favoring influence upon root formation in the more basally situated parts of the stem is one of the factors determining the polar character of regeneration.     

Abscisic acid inhibits shoot elongation of Scirpus mucronatus

The relationships between free ABA levels and shoot elongation were investigated in shoots of Scirpus mucronatus L. Under submergence, shoot elongation increased but free ABA levels decreased. The extent of the increase in length and the decrease in free ABA in submerged shoots increased with the increase of water depth. When the shoots were transferred to air after 12 days of submergence, they ceased to elongate and the free ABA levels recovered to the values of air-grown shoots. ABA, at concentrations from 1 μ M to 1 m M , inhibited the submergence-induced shoot elongation. In ambient air, fluridone, an inhibitor of ABA biosynthesis, at 7 μ M decreased the free ABA levels in shoots but increased shoot elongation. The effects could be reversed by 10 μ M ABA. These results indicate that ABA is an internal inhibitor of shoot growth in Scirpus .     

Stock-plant etiolation causes drifts in total soluble sugars and anthraquinones, and promotes adventitious root formation in teak (Tectona grandis L. f.) coppice shoots

The effects of stock-plant etiolation on coppice-shoot growth, drifts in total soluble sugars and anthraquinones (AQs; C₁₄H₈O₂), and rooting potentiality of shoot cuttings were examined in Tectona grandis L. f. (clone FG1). When seedlings were one-year-old, they were coppiced and maintained in the dark for etiolation, with a parallel set kept under natural light in an open environment. Coppice shoots were made into single-node leafy cuttings (SNCs), which were cultured under intermittent mist for rooting. These SNCs were treated with different concentrations of NAA (0, 2000 and 3000 mg l⁻¹). Etiolation significantly increased the coppice-shoot length, internode length, number of coppice shoots, number of leaves, number of nodes and total soluble sugars. The HPTLC analysis showed qualitative and quantitative differences in AQs in coppice shoots obtained from etiolated and non-etiolated stock plants. The study showed that AQs could be used as a marker for maturity and juvenility in teak. Stock-plant etiolation caused a significant increase in percent rooting and sprouting, shoot length, number of shoots and number of leaves per SNC, but a decrease in callusing at the base of the SNC. NAA at 2000 and 3000 mg l⁻¹ had inhibitory effects on rooting and sprouting of SNCs. The result showed that stock-plant etiolation fostered rooting by rejuvenating the coppice shoots.     

Why does the bud-galling sawfly,Euura mucronata,attack long shoots?

Summary The bud-galling sawfly, Euura mucronata, attacked longer shoot length classes on its host, Salix cinerea, more frequently than shorter shoots. Shoot length accounted for 76 to 93 percent of the variance in number of galls per 100 shoots in three habitats: forest, watermeadow, and lakeside. The reasons for this pattern were addressed with studies on shoot length in relation to: 1. Number of resources (buds) per shoot; 2. Success in establishment of larvae in galls; 3. Gall size and resources per gall; and 4. Survival of larvae after establishment as influenced by plant resistance and natural enemy attack. The most important factors proved to be success in establishment of larvae, with percent of variance accounted for ranging from 57 to 77 percent in three of four sites where relationships were significant, and survival after establishment of larvae, with variance accounted for ranging from 40 to 54 percent in the same three sites. The pattern of survival was dictated by plant resistance and not by natural enemies. These two additive factors resulted in a general relationship across all sites of increasing emergence of fully developed larvae per cohort as shoot length increased, accounting for 78 percent of the variance. These adaptive advantages to attacking longer shoots are sufficient to account for the pattern of increased probability of shoots being attacked as they increase in length.     

Differential Responses of Buds along the Shoot to Factors Involved in Apical Dominance

Intact and decapitated 6-node shoots of Hygrophila sp. weregrown aseptically immersed in liquid half-strength Knop's solutionwith microelements and 2% (w/v) sucrose (control medium), andin medium with 0.1 mg l^–1 benzyladenine (BA). In intactshoots grown in control medium apical dominance suppressed outgrowthof the lateral buds; in decapitated shoots buds grew out atseveral of the most apical nodes, increasing in size acropetally.There was a lag in outgrowth of the bud at the most apical node,attributable to its initially smaller size. Lateral shoots grewout first at basal nodes of intact shoots in BA medium, decreasingin size acropetally; in decapitated shoots in BA medium lateralshoots of approximately equal size grew out at all nodes. Differentialeffects of decapitation and cytokinin treatment on lateral shootoutgrowth along the shoot could be interpreted by postulatinga basipetally decreasing gradient of endogenous auxin concentrationin the intact shoot. Application of 20 mg l^–1 indoleaceticacid (IAA) in agar to decapitated shoots completely preventedbud outgrowth for at least 7 d in control medium, inhibitingit thereafter, and inhibited bud outgrowth in BA medium, thussupporting the hypothesis. Comparison of lateral shoot outgrowthin whole decapitated shoots and severed decapitated shoots (isolatednodes) lent no support to the alternative hypothesis that theremight be an acropetally decreasing concentration gradient ofa bud-promoting substance in the intact shoot, and demonstratedmuch greater lateral shoot growth in isolated nodes. The resultsemphasize important correlative relationships between the partsof a shoot with several nodes.     

Willow regrowth after galling increases bud production through increased shoot survival

Insect herbivory can negatively or positively affect plant performance. We examined how a stem gall midge Rabdophaga rigidae affects the survival, growth, and bud production of current year shoots of the willow Salix eriocarpa. In mid-May, the gall midge initiates stem galls on the apical regions of shoots. The following spring, galled shoots had thicker basal diameters and more lateral shoots than ungalled shoots. Although galled shoots were on average 1.6 times longer than ungalled shoots, there were no significant differences in shoot length or in the numbers of reproductive, vegetative, and dormant buds per shoot. However, the subsequent survival of galled shoots was significantly higher than that of ungalled shoots, probably because of the thicker basal diameter. This increased shoot survival resulted in approximately two times greater reproductive, vegetative, and dormant bud production on galled shoots compared with ungalled shoots in the following spring. These results suggest that the willow regrowth induced by galling can lead to an increase in bud production through increased shoot survival.     

A rapid in vitro propagation protocol for Piper barberi Gamble, a critically endangered plant

Summary A protocol is described for rapid multiplication of Piper barberi Gamble (Piperaceae) through shoot tip and nodal explant cultures. Nodal explants with a single axillary meristem showed three times better response with respect to shoot proliferation when compared to shoot tip explants. The best shoot proliferation response of nodal explants was observed with a cytokinin combination of N₆-benzyladenine (4.43 μM) and kinetin (2.32 μM), with 88% bud break. The number of shoot initials (2.4) produced per nodal explant was twice the number of shoot initials (1.2) per shoot tip. An average of 6.9±0.58 adventitious shoots were observed from the proximal end of the internodal explants on Mursashige and Skoog (1962) (Ms) basal medium supplemented with N₆-benzyladenine (2.22 μM) and kinetin (0.46 μM). A multiplication rate of 82 shoots per explant could be achieved after 9 wk of subculturing. The in vitro shoots were rooted on one-half and one-quarter MS basal medium. The shoots rooted on one-quarter MS in the dark produced eight roots with an average root length of 3.36 cm and 98% survival. These plants were transferred to the field with a survival rate of 75%.     

A two-step procedure for adventitious shoot regeneration on excised leaves of lowbush blueberry

An efficient system to regenerate shoots on excised leaves of greenhouse-grown wild lowbush blueberry (Vaccinium angustifolium Ait.) was developed in vitro. The effect of thidiazuron (TDZ) on adventitious bud and shoot formation from apical, medial, and basal segments of the leaves was tested. Leaf cultures produced multiple buds and shoots with or without an intermediary callus phase on 2.3–4.5 μM TDZ within 6 wk of culture initiation. The greatest shoot regeneration came from young expanding basal leaf segments positioned with the adaxial side touching the culture medium and maintained for 2 wk in darkness. Callus development and shoot regeneration depended not only on the polarity of the explants but also on the genotype of the clone that supplied the explant material. TDZ-initiated cultures were transferred to medium containing 2.3–4.6 μM zeatin and produced usable shoots after one additional subculture. Elongated shoots were dipped in 39.4 mM indole-3-butyric acid powder and planted on a peat:perlite soilless medium at a ratio of 3:2 (v/v), which yielded an 80–90% rooting efficiency. The plantlets were acclimatized and eventually established in the greenhouse with 75–85% survival.     

In vitro induction of multiple shoots and plant regeneration in cotton (Gossypium hirsutum L.)

Induction of multiple shoots in cotton (Gossypium hirsutum L. cv. Anjali-LRK 516) has been achieved with cotyledonary nodes devoid of cotyledons and apical meristems. Explants from 35-day-old seedlings yielded the maximum number of shoots (4.7 shoots/explant) using Murashige and Skoog (MS) basal medium supplemented with 6-benzylaminopurine and kinetin (2.5 mg/1 each). Explants from 35-day-old seedlings raised in glass bottles produced a higher number of multiple shoots (8.3 shoots/explant) than those grown in glass tubes and cultured on the same shoot induction medium. Elongation of multiple shoots was obtained on liquid or agar MS basal medium without phytohormones. In vitro shoots were rooted on half-strength agar-solidified MS basal medium or with 0.05 or 0.1 mg/1 naphthaleneacetic acid. Hardening and survival of tissue culture plantlets was 95% under greenhouse conditions.Abbreviations BAP 6-Benzylaminopurine - GA₃ Gibberellic acid - MS Murashige and Skoog medium - NAA -Napthaleneacetic acid     

Branching responses in Silphium integrifolium (Asteraceae) following mechanical or gall damage to apical meristems and neighbor removal

Branching in plants increases plant access to light and provides pathways for regrowth following damage or loss of the apical meristem. We conducted two experiments in an eastern Kansas tallgrass prairie to determine how apical meristem loss (by clipping), apical meristem damage (by insect galling), and increased light availability affected growth, reproduction, and branching in Silphium integrifolium (Asteraceae). The first experiment compared clipping with galling. Clipping increased axillary shoot numbers, while galling increased axillary shoot lengths, reflecting different allocation responses among damage types and inhibition of branching by galls. However, total capitulum production was less in all gall/clip treatments than in intact shoots. The second experiment compared clipping with mowing the surrounding vegetation to increase light availability. Mowing increased total leaf, total capitulum, and axillary shoot length and axillary capitulum production in clipped and unclipped plants and in large vs. small shoots. The presence of the neighboring canopy, not of an intact apical meristem, was therefore the stronger limitation on leaf and capitulum production. These experiments suggest that damage and light competition affected both branching frequency and the partitioning of resources among shoots, branches, and leaves. Because Silphium's growth form is widespread, similar responses may occur in other grassland forbs.