Periods of organogenesis in mono- and bicyclic annual shoots of Juglans regia L. (Juglandaceae)

The organogenetic cycle of shoots on main branches of 4-year-old Juglans regia trees was studied. Mono- and bicyclic floriferous and vegetative annual shoots were analysed. Five parent annual shoot types were sampled between October 1992 and August 1993. Organogenesis of summer growth units was monitored between 16 Jun. and 3 Aug. 1993. Variations over time in the number of nodes, cataphylls and embryonic green leaves of terminal buds were studied. The number of nodes of parent shoot buds was compared with the number of nodes of shoots derived from parent shoot buds. The spring growth units of mono- and bicyclic shoots consist exclusively of preformed leaves which were differentiated, respectively, during the spring flush of growth (mid-April until mid-May) or the summer flush of growth (mid-June until early August) in the previous growing season. Thus, winter buds may consist of flower and leaf primordia differentiated in two different periods during annual shoot extension. The summer growth units of bicyclic shoots consist of preformed leaves that were differentiated in spring buds during the spring flush of growth in the current growing season. Bud morphology is compared between spring and summer shoots.     

Application of architectural analysis and AMAPmod methodology to study dwarfing phenomenon: the branch structure of 'Royal Gala' apple grafted on dwarfing and non-dwarfing rootstock/interstock combinations

Architectural analysis was applied to study branch development of 'Royal Gala' apple trees grafted with dwarfing and non-dwarfing rootstock/interstock combinations, which had been chosen to produce trees with a wide range of vigour. Using AMAPmod methodology, the structure of 3-year-old branches was described at four levels of representation: branch; annual shoot; growth unit; and node. Three types of growth units were distinguished: extension growth unit (vegetative unit with internode extension); vegetative spur with minimal internode extension; and fruiting spur or bourse. The aim of the analysis was to describe exactly how the rootstock/interstock combinations affected the structure building process. The number of extension growth units, vegetative spurs and fruiting spurs per annual shoot changed over the years, but this was not affected by rootstock/interstock combination. Compared with MM.106 rootstock, M.9 rootstock reduced the number of nodes per extension growth unit. In most cases, rootstock/interstock combination had no effect on the linear relationship between extension growth unit length and node number (R(2) = 0.88). Average internode length depended on unit node number, with internodes being shorter for units with fewer nodes. Thus the difference in apple branch size induced by the rootstock/interstock combinations was mainly due to a reduction in the length and number of neoformed nodes produced on extension growth units. As percentage budbreak of axillary buds on extension growth units was not affected by rootstock/interstock combination, differences in numbers of axillary annual shoots per branch were entirely due to differences in the total numbers of nodes extended during the previous year.     

Apple dwarfing rootstocks and interstocks affect the type of growth units produced during the annual growth cycle: precocious transition to flowering affects the composition and vigour of annual shoots

Background and Aims: Precocious flowering in apple trees is often associated witha smaller tree size. The hypothesis was tested that floral evocationin axillary buds, induced by dwarfing rootstocks, reduces thevigour of annual shoots developing from these buds comparedwith shoots developing from vegetative buds. Methods: The experimental system provided a wide range of possible treevigour using ‘Royal Gala’ scions and M.9 (dwarfing)and MM.106 (non-dwarfing) as rootstocks and interstocks. Second-yearannual shoots were divided into growth units corresponding toperiods (flushes) of growth namely, vegetative spur, extensiongrowth unit, uninterrupted growth unit, floral growth unit (bourse)and extended bourse. The differences between the floral andvegetative shoots were quantified by the constituent growthunits produced. Key Results: The dwarfing influence was expressed, firstly, in reduced proportionsof shoots that contained at least one extension growth unitand secondly, in reduced proportions of bicyclic shoots (containingtwo extension growth units) and shoots with an uninterruptedgrowth unit. In treatments where floral shoots were present,they were markedly less vigorous than vegetative shoots withrespect to both measures. In treatments with M.9 rootstock,vegetative and floral shoots produced on average 0·52and 0·17 extension growth units, compared with 0·77extension growth units per shoot in the MM.106 rootstock treatment.Remarkably, the number of nodes per extension growth unit wasnot affected by the rootstock/interstock treatments. Conclusions: These results showed that rootstocks/interstocks affect thetype of growth units produced during the annual growth cycle,reducing the number of extension growth units, thus affectingthe composition and vigour of annual shoots. This effect isparticularly amplified by the transition to flowering inducedby dwarfing rootstocks. The division of annual shoot into growthunits will also be useful for measuring and modelling effectsof age on apple tree architecture.     

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.     

Growth Dynamics and Morphology of Annual Shoots, According to their Architectural Position, in Young Cedrus atlantica (Endl.) Manetti ex Carriere (Pinaceae)

The influence of shoot architectural position on growth andbranching pattern of young Cedrus atlantica (Endl.) Manettiex Carrière trees were studied. Extension growth andtype of axillary products (lateral bud, sylleptic short or longshoots) of annual shoots of increasing branching order (mainstem, branches and branchlets) were recorded weekly during the1993 growing season. Annual final shoot length, duration ofextension, and maximum extension rate decreased with increasingbranching order. Sylleptic axillary shoots occurred only onannual shoots of the main stem and branches and were producedwhen extension rate was at its highest. Differences in growthrate and final length of annual shoots, according to their architecturalposition, were related to differences in the total number anddiversity of types of sylleptic axillary shoots produced. Itis suggested that types and numbers of sylleptic axillary shootsproduced are linked with threshold values for both final lengthand extension rate of the parent shoot. Copyright 1999 Annalsof Botany Company Atlas cedar, extension growth, sylleptic branching, tree architecture, morphology.     

Differences in proleptic and epicormic shoot structures in relation to water deficit and growth rate in almond trees (Prunus dulcis)

Background and Aims

Shoot characteristics differ depending on the meristem tissue that they originate from and environmental conditions during their development. This study focused on the effects of plant water status on axillary meristem fate and flowering patterns along proleptic and epicormic shoots, as well as on shoot growth rates on ‘Nonpareil’ almond trees (Prunus dulcis). The aims were (1) to characterize the structural differences between proleptic and epicormic shoots, (2) to determine whether water deficits modify shoot structures differently depending on shoot type, and (3) to determine whether shoot structures are related to shoot growth rates.

Methods

A hidden semi-Markov model of the axillary meristem fate and number of flower buds per node was built for two shoot types growing on trees exposed to three plant water status treatments. The models segmented observed shoots into successive homogeneous zones, which were compared between treatments. Shoot growth rates were calculated from shoot extension measurements made during the growing season.

Key Results

Proleptic shoots had seven successive homogeneous zones while epicormic shoots had five zones. Shoot structures were associated with changes in growth rate over the season. Water deficit (1) affected the occurrence and lengths of the first zones of proleptic shoots, but only the occurrence of the third zone was reduced in epicormic shoots; (2) had a minor effect on zone flowering patterns and did not modify shoot or zone composition of axillary meristem fates; and (3) reduced growth rates, although patterns over the season were similar among treatments.

Conclusions

Two meristem types, with different latency durations, produced shoots with different growth rates and distinct structures. Differences between shoot type structure responses to water deficit appeared to reflect their ontogenetic characteristics and/or resource availability for their development. Tree water deficit appeared to stimulate a more rapid progression through ontogenetic states.     

Relative importance of nodal roots and apical buds in the control of branching in Trifolium repens L.

Clonal species are characterised by having a growth form in which roots and shoots originate from the same meristem so that adventitious nodal roots form close to the terminal apical bud of stems. The nature of the relationship between nodal roots and axillary bud growth was investigated in three manipulative experiments on cuttings of a single genotype of Trifolium repens. In the absence of locally positioned nodal roots axillary bud development within the apical bud proceeded normally until it slowed once the subtending leaf had matured to be the second expanded leaf on the stem. Excision of apical tissues indicated that while there was no apical dominance apparent within fully rooted stems and very little in stems with 15 or more unrooted nodes, the outgrowth of the two most distal axillary buds was stimulated by decapitation in stems with intermediate numbers of unrooted nodes. Excision of the basal branches from stems growing without local nodal roots markedly increased the length and/or number of leaves on 14 distally positioned branches. The presence of basal branches therefore prevented the translocation of root-supplied resources (nutrients, water, phytohormones) to the more distally located nodes and this caused the retardation in the outgrowth of their axillary buds. Based on all three experiments we conclude that the primary control of bud outgrowth is exerted by roots via the acropetal transport of root-supplied resources necessary for axillary bud outgrowth and that apical dominance plays a very minor role in the regulation of axillary bud outgrowth in T. repens.     

PHENOTYPIC PLASTICITY OF BRANCHING PATTERN IN SUGAR MAPLE (ACER SACCHARUM)

Branching patterns of leader shoots and lower branches were compared in 15 sugar maple trees. The number of first-order branches per second-order branch (R_1:2) differed very significantly between leader shoots and lower branches. Values of R_1:2 averaged 6.47 for leaders and 4.06 for lower branches. Stepwise bifurcation ratios between successive higher orders did not differ significantly. The difference in R_1:2 between leader shoots and lower branches within individual trees is the result of developmental-phenotypic interactions, such that the number of leaf-bearing shoots per support shoot decreases over time. These results indicate that branching patterns within individual trees are not stationary, as has commonly been claimed.     

AXR1 acts after lateral bud formation to inhibit lateral bud growth in Arabidopsis

The AXR1 gene of Arabidopsis is required for many auxin responses. The highly branched shoot phenotype of mature axr1 mutant plants has been taken as genetic evidence for a role of auxin in the control of shoot branching. We compared the development of lateral shoots in wild-type Columbia and axr1-12 plants. In the wild type, the pattern of lateral shoot development depends on the developmental stage of the plant. During prolonged vegetative growth, axillary shoots arise and develop in a basal-apical sequence. After floral transition, axillary shoots arise rapidly along the primary shoot axis and grow out to form lateral inflorescences in an apical-basal sequence. For both patterns, the axr1 mutation does not affect the timing of axillary meristem formation; however, subsequent lateral shoot development proceeds more rapidly in axr1 plants. The outgrowth of lateral inflorescences from excised cauline nodes of wild-type plants is inhibited by apical auxin. axr1-12 nodes are resistant to this inhibition. These results provide evidence for common control of axillary growth in both patterns, and suggest a role for auxin during the late stages of axillary shoot development following the formation of the axillary bud and several axillary leaf primordia.     

Flowering and fruiting phenology in the apomictic blackberry, Rubus nessensis (Rosaceae)

Five stands of an apomictic blackberry, Rubus nessensis , were studied. Biennial shoots flower and fruit in their second year on lateral branches developed from axillary buds. Most premature death of reproductive units occurs by withering of entire laterals, presumably due to damage by frost during the preceeding autumn or winter. Health status of the floricane, assessed as percentage live axillary buds in the spring, was positively correlated with the reproduction parameters. Shoot height as well, is positively correlated with number of flower buds and flowers produced on each lateral but, however, negatively correlated with number of berries, probably because tall shoots tend to suffer more damage by frost which seriously afflicts the vascular tissues of the laterals.
The position of the axillary buds on the shoot affects survival and degree of fertility, those situated medially greatly surpassing apical and basal ones.     

Analysis of Branching in Spring-sown White Lupins (Lupinus albusL.): The Significance of the Number of Axillary Buds

Plant density and sowing date were shown to affect branchingin spring-sown white lupin (Lupinus albusL.), but the responsevaried among environments. The patterns of primary and secondarybranching in the cv. Lublanc were studied as a function of boththe number of axillary buds and the plant growth rate. Fieldexperiments that used a wide range of sowing dates and plantdensities to alter plant architecture were conducted over 5years, and these were supplemented with data from additionalglasshouse and growth cabinet experiments. The number of axillary buds on the main stem or primary branches,which determined the potential number of branches, increasedlinearly with the number of nodes. In situations where all axillarybuds did not produce branches, it was found that the numberof primary and secondary branches produced was related to theplant growth rate at the beginning of branch elongation. Knowledgeof the number of axillary buds improved the analysis of theinteraction between cultural practices and environmental conditionson plant architecture. The variability of branching potentialamong genotypes was discussed. Lupinus albus; branches; axillary buds; growth; sowing date; plant density     

Interactive effects of herbivory and sand burial on growth of a tropical dune plant, Ipomoea pes-caprae

1. This study examined the effects of insect herbivory, sand burial, and the interactive effects of these factors, on the growth of beach morning glory, Ipomoea pes-caprae, a common tropical dune plant. Levels of herbivory and sand burial were manipulated on individual shoots, and effects on stem growth, leaf production, and production of adventitious roots and axillary branches by nodes were examined.
2. Sand burial had a significant positive effect on the production of roots, but did not affect growth in stem length or leaf production.
3. Effects of herbivory were consistently negative, and persisted for 6 weeks after the herbivore damage was incurred. Stem growth rates and leaf production decreased and the production of bare nodes (with no roots or branches) and mortality of apical meristems increased.
4. Interactive effects of herbivory and sand burial influenced both leaf production and root production. In the case of root production, the effects of herbivory in decreasing the proportion of nodes that produced roots occurred only in the presence of burial. In contrast, burial masked the negative effects of herbivory on leaf production.     

Meristem growth dynamics and branching patterns in the Cladoniaceae

Branching patterns in the lichen family Cladoniaceae are varied and taxonomically important. Branching occurs on the podetium, the erect secondary thallus that characterizes most species in the Cladoniaceae, and is influenced by growth dynamics of the fungal meristem tissue at the apex of the podetium. Branching is primarily the result of meristem divisions, and branching patterns are modified by meristem enlargement, deformation, and torsion. Branching processes are conserved, and early branch ontogeny provides information from which to determine relationships in the Cladoniaceae. Branching is characterized by two major patterns. In one pattern, branches arise from the relatively late divisions of a large meristem (≥100 μm in diameter), whose shape changes during ontogeny. In a second pattern, branches arise from small meristems (<100 μm in diameter), which split early in ontogeny but whose shape does not change. The trend toward reduced meristems that split early in ontogeny is seen as an evolutionary advance in the Cladoniaceae. Some "small meristem" species retain aspects of the "large meristem" habit in early ontogeny, and this provides a clue to their relationships. Patterns of meristem growth dynamics provide a basis for interpreting phylogeny in mycobionts of the Cladoniaceae. Meristem activities in four genera of the Cladoniaceae were compared in order to determine trends in growth dynamics within the family.     

A study of the autotropic straightening reaction of a shoot previously curved during geotropism

Abstract The growth of marked zones on horizontal sunflower and maize shoots was measured over long periods. Subsequent to a period of geocurvature, a period of autotropic straightening occurred in most zones of these shoots. The patterns of differential growth causing this straightening were determined. The straightening appeared in the more apical zones initially and subsequently in the basal zones. The magnitude of the differential growth causing the straightening in any zone was similar to that which caused geocurvature in the same zone. It was found that autotropic straightening became evident in some zones before those zones had reached a vertical orientation thus it is suggested that gravity is not directly involved in the initiation of autotropism.     

Significance of the simultaneous growth of vegetative and reproductive organs in the prostrate annual Chamaesyce maculata (L.) Small (Euphorbiaceae)

To elucidate the significance of the simultaneous growth of vegetative and reproductive organs in the prostrate annual Chamaesyce maculata (L.) Small (Euphorbiaceae) from the standpoint of meristem allocation, we investigated plant architecture, meristem allocation, and the spatial and temporal patterns in vegetative growth and reproduction in the reproductive stage. The numbers of secondary and tertiary shoots successively increased by branching in the reproductive stage, and the sum of shoot length was greater in secondary shoots than in primary shoots. The specific shoot length (shoot length per shoot biomass) was greater in lateral shoots than in primary shoots, indicating efficient lateral shoot elongation. The internode length was shorter in secondary shoots than in primary shoots, increasing the number of nodes per shoot length in secondary shoots. Many nodes on a shoot generated two meristems, one of which committed to a flower and one to a lateral shoot. The number of reproductive meristems was greatest in tertiary shoots, and 96% of total reproductive meristems on shoots were generated in lateral shoots. On almost all nodes, the reproductive meristem developed into a flower, and 95–98% of the flowers produced a fruit. Therefore, vegetative growth by branching in the reproductive stage contributed to the increase in reproductive outputs. From the standpoint of meristem allocation, the simultaneous growth of vegetative and reproductive organs in prostrate plant species might be important for increasing the number of growth and reproductive meristems, resulting in the increase in reproductive outputs.     

Floral-stimulus activity in tobacco stem pieces

The growth patterns of axillary buds of dayneutral tobacco (Nicotiana tabacum L. cv. Wisconsin 38) plants were assessed by using expiants of single buds attached to leafless stem cuttings and allowing the buds to grow to flowering without additional manipulation. Buds located 5, 10 and 15 nodes below the inflorescence were employed. For a given bud position, when a cutting had few internodes the growth pattern of a bud tended to fall into one of two groups: buds that produced few-noded shoots and buds that produced many-noded shoots. For example, in a group of 13 cuttings composed of bud 5 with 2 associated internodes, 11 buds produced 14.2 nodes (range, 11–17) and 2 buds produced 32.0 nodes (range, 30–34). As the number of internodes on the cutting increased, the number of buds producing few-noded shoots increased and the number of nodes produced decreased (e.g. in contrast to the data above, all 5th buds with 6 internodes produced 12.8 nodes; range 11–15). When cuttings from the 3 positions had the same number of internodes, the more apical cuttings had buds that produced fewer nodes (e.g. for cuttings with 6 internodes all 5th buds produced 12.8 nodes, all 10th buds produced 15.5 nodes and 85% of 15th buds produced few-noded shoots with 19.3 nodes). The number of nodes produced by a bud was a function of the original position of the stem piece and not the original position of the bud. That is, bud 5 associated with the 6 internodes below it produced 12.8 nodes and bud 10 associated with essentially the same 6 internodes (i.e. the 6 above it) produced 12.9 nodes while bud 10 associated with the 6 internodes below it produced 15.5 nodes. Thus, the number of nodes produced by a bud was dependent upon the original main-axis position of the cutting as well as the number of internodes on the cutting. Buds forced to grow out in situ on main axes devoid of leaves produced substantially more nodes than similar buds on cuttings. Buds isolated without associated internodes produced many-noded plants with a number of nodes similar to that of plants grown from seed. The simplest interpretation of these data is that stem pieces contain floral-stimulus activity and that this activity is present in a gradient with the highest activity being located in the apical part of the stem.We thank Susan Smith and Harry Roy (Rensselaer) for comments, and the National Science Foundation for financial support (IBN-9003739 to C.N.M.).     

Axillary shoot proliferation and growth of Populus deltoides shoot cultures

Shoot cultures of four genotypes of Populus deltoides Bartr. ex Marsh. were established from adventitious shoots regenerated from internodal stem explants. Stable shoot cultures for all four genotypes were maintained in a continuous culture regime for over one year. The stable shoot cultures were used as explants to investigate the effects of zeatin concentration and genotype on axillary shoot production and growth. The concentration of zeatin significantly affected the production of axillary shoots, with 1.0 mgL^–1 zeatin producing the greatest number of shoots (31.0 shoots per culture vessel) while 0.25 mgL^–1 zeatin produced the greatest growth (5.9 mg per axillary shoot) when measured by dry weight accumulation per shoot. Genotypic differences were significant in the production and growth of axillary shoots.Abbreviations DKW Driver and Kuniyuki Walnut medium - PAR Photosynthetically Active Radiation Journal Series No. 9111, Agricultural Research Division, University of Nebraska     

The origin, initiation and development of axillary shoot meristems in Lotus japonicus

BACKGROUND AND AIMS: Lotus japonicus 'Gifu' develops multiple axillary shoots in the cotyledonary node region throughout the growth of the plant. The origin, initiation and development of these axillary meristems were investigated. METHODS: Morphological, histological and mRNA in situ analyses were done to characterize the ontogeny of cotyledonary axillary shoot meristems in Lotus. Morphological characterization of a putative Lotus shoot branching mutant (super-accessory branches) sac, is presented. KEY RESULTS: By using expression of an L. japonicus STM-like gene as a marker for meristematic tissues, it was demonstrated that groups of cells maintained in the meristematic state at the cotyledonary axil region coincide with the sites where additional axillary meristems (accessory meristems) form. A Lotus shoot branching mutant, sac, is a putative Lotus branching mutant characterized by increased proliferation of accessory shoots in all leaf axils including the cotyledons. CONCLUSION: In Lotus, axillary shoot meristems continually develop at the cotyledonary node region throughout the growth of the plant. These cotyledonary primary and accessory axillaries arise from the position of a meristematic zone of tissue at the cotyledonary node axil region.