The Shoot Apex and its Dichotomous Branching in the Nypa Palm

Seedlings of the palm Nypa fruticans van Wurmb are viviparous,the plumule becoming exserted before the fruit is ripe and possiblyassisting in fruit detachment. Established seedlings have horizontalaxes, this growth orientation being maintained throughout thelife of the palm which may be described as ‘rhizomatous’.Inflorescences are axillary in adult plants, but their distributionis irregular. The shoot apex is small and very asymmetricalsince it is more or less displaced into a lateral position byprogressive enlargement of each leaf primordium during thisleaf's first plastochrone. The plastochrone interval is apparentlya long one so that leaves have a wide developmental gap betweenthem. This results in a leaf base which is more or less cylindricalbut with a groove to accommodate the next youngest leaf. All the available evidence suggests that vegetative branchingis by equal dichotomy of the shoot apex at wide intervals. Thedichotomy is marked superficially by a leaf, enclosing the twonew shoots, which has two grooves. The twin shoots are insertedin the lateral, not the dorsiventral plane of this enclosingleaf. The daughter shoots are always at precisely the same stageof development and they always have mirror-image symmetry withrelation to each other, the phyllotaxis of the parent shootbeing maintained without obvious interruption. There is no anatomicalevidence for abortion of the original apex and its replacementby two new ones. The vascular system is divided equally betweeneach daughter shoot without interruption, suggesting stronglythat there is continuity of growth from the undivided to thedivided condition.     

PHYLLOTAXIS AND VASCULAR ORGANIZATION OF THE CARPELS IN MICHELIA FUSCATA

Tucker , Shirley C. (U. Minnesota, Minneapolis.) Phyllotaxis and vascular organization of the carpels in Michelia fuscata. Amer. Jour. Bot. 48(1): 60–71. Illus. 1961.—Phyllotaxis pattern and vascular organization are closely related in the floral receptacle of Michelia fuscata (Magnoliaceae). The carpels arise in a spiral or helix. They are initiated alternately along each of 7, 8 or 10 helical parastichies according to a complex repetitive sequence. The pattern of the dorsal carpellary trace fusions is orderly for each of the 10 flowers investigated. The dorsal carpellary traces in each parastichy diverge from the same vascular sympodium. Among flowers one finds differing numbers of parastichies, differing angles of divergence, and varying sequences of parastichies which reflect the order of carpel initiation. The angle of divergence, although consistent for any 1 parastichy in a flower, can vary greatly between parastichies. The nature and importance of the organizers which determine appendage position at the apical meristem are considered. Changes in apical size, configuration, and activity are shown to be related to phyllotaxis.     

The Morphogenesis of Apple Buds: IV. The Effect of Fruit

The course of development of spur buds on fruiting trees of‘biennial’ and ‘regular’ apple varietieswas followed throughout the season by dissection of successivesamples of buds and the effect on bud development of de-fruitingtrees at two different times in the season was recorded. In the biennially fruiting variety Miller's Seedling, the patternof bud development on fruiting trees was very similar to thatin buds on non-fruiting trees which failed to form flowers followingdefoliation, as were the number and sizes of leaves precedingthe bud in each instance. The failure to form flowers was associatedwith the occurrence of an 18-day plastochrone in buds, and itwas concluded that on fruiting trees it was not due to a competitiveeffect of the fruit for nutrients. This long plastochrone was not found in buds of de-blossomedtrees of Miller's Seedling, and in trees de-fruited later inthe season the buds immediately broke into a new flush of growthand at the same time the plastochrone was shortened. These resultssuggest that the long plastochrone was due to the inhibitoryeffect of fruit on the older primordia of the bud, an effectwhich did not occur until after the resting buds had begun toform. A phase with an 18-day plastochrone was also found inbuds of another biennial variety, Laxton's Superb, but not inthose of the regularly fruiting variety Sunset. Developing fruitlets of biennial varieties caused bud-scalesto form sooner and hastened their rate of development, possiblydue to changes induced in the levels of a gibberellin-like apexfactor in the buds. The rate of increase in number of bud-scalesin the bud appeared to depend upon the extent to which the budwas affected by the primary leaves of the flower cluster andthose of other clusters.     

A contact pressure model for semi-decussate and related phyllotaxis

Semi-decussate phyllotaxis, in which leaves arise singly and the divergence angles between successive pairs of leaves alternate between approximately 90° and approximately 180°, is accounted for by a contact pressure model. It is assumed that leaf primordia are initiated at a divergence angle close to the Fibonacci angle of 137·5°, that the primordia move under contact pressure, and that when a primordium first experiences contact pressure all other primordia are fixed. Extensions of the model account for: psuedodecussate phyllotaxis, where the leaves appear to arise in pairs; semi-tricussate and pseudo-tricussate phyllotaxis, where the leaves are arranged in, respectively, dissolved or apparent trimerous whorls; and phyllotaxis of the 1,3 series, where the divergence angle is about 100°. The compatibility of the model with current theories of Fibonacci phyllotaxis is discussed.     

Allometric relations in plant growth

Allometric relations Y = aX^b are shown to exist in phyllotaxis, under the form r = k log R = p ()(m + n) ^–2 , where r = Y is the normalized internode distance in the cylindrical representation of phyllotaxis and R is the plastochrone ratio in the centric representation, p()=a is a constant for every angle of intersection of the opposed parastichies of the visible pair (m, n), for every m and n, and for all possible limit divergence angles corresponding to the Fibonacci-type sequences..., m–n, n, m, m + n=X, 2m + n, ..., and where b=–2. Richards' phyllotaxis index will be deduced.This work was supported by the Natural Science and Engineering research Council Canada, grant A6240     

Leaf phyllotaxis: Does it really affect light capture?

The intriguing mathematical properties of leaf phyllotaxis still attract scientific attention after centuries of research. Phyllotaxis, and in particular the divergence angle between successive leaves, have been frequently interpreted in terms of maximization of light capture, although certain model simulations of light capture by vertical shoots revealed minor effects of phyllotaxis in comparison with the effect of other morphological features of the plant. However, these simulations assumed a number of simplifications, did not take into account diffuse light, and were not based on real plants with their natural range of morphological variation. This study was aimed at filling these gaps by examining the influence on light harvesting of shoot architecture and divergence angle in four species with spiral phyllotaxis (Quercus ilex, Arbutus unedo, Heteromeles arbutifolia and Daphne gnidium) with a realistic 3-D model (Y-plant). A wide range of divergence angles (from 100° to 154°) was observed within each species, with 144° being the most frequent one. These different divergence angles rendered very different vertical projections of the shoot due to contrasting patterns of leaf overlap as seen from above, but they rendered indistinguishable light interception efficiencies (Ea). Setting the leaves with an opposite-decussate phyllotaxis led, however, to a 40–50% decrease of Ea. The interplay of internode length, leaf size and shape, and leaf elevation angle led to significant species differences in Ea. Thus, only particular phyllotaxis (e.g., decussate) might be functionally inefficient under certain combinations of the various morphological variables that influence light capture of a shoot. This revised version was published online in June 2006 with corrections to the Cover Date.     

Comparative Studies of Seed Germination of two Newly Isolated Lines of Grand Rapids Lettuce

Two lines of Grand Rapids lettuce were isolated by self-pollinationof single plants. Seeds of one line, designated ‘A’,germinate in the dark at 25 °C. Seeds of the second line,designated ‘B’ germinate in a manner typical oftheir variety. The lines differ in sensitivity to red (R) andfar-red (FR) light, in the ability of their seeds to overcomeresistance of an external osmotically active medium, in theresponse of mature seeds to abscisic acid, and in the responseof young seeds to gibberellin and benzyl adenine. ‘A’seeds germinate faster than ‘B’ seeds in the darkor after R or FR irradiation.     

A Quantitative Study of Shoot Development in Vicia faba: III. The Dry Weights of the Plumular Internodes

The development of the bean shoot has been studied by cuttinga piece out of each internode and determining the dry weightper unit stem length, the ‘linear density’ of theintemode. The measurements of this quantity are used in the mathematicalanalysis of physiological age. It is shown that there is a drasticreduction in relative growth rate of the stem during the unfoldingof the ninth leaf and that there is evidence of an unknown factoraffecting many of the shoot-growth processes with a maximumintensity in the fifth internode. The bearing of these factson the conception of the plastochrone as a unit of shoot growthis discussed.     

PHOTOPERIOD INDUCED CHANGE IN PHYLLOTAXIS IN XANTHIUM

Photoperiodic floral induction in Xanthium, achieved by subjecting the plants to two long nights, is accompanied by a transient change of the phyllotaxis from the (2, 3) contact parastichy pattern of vegetative plants, to a (3, 5) pattern during the transition. To specify the phyllotaxis, two parameters were estimated from transverse sections of apical buds of control and treated plants: the divergence angle, α, and the plastochron ratio, a. The plastochron ratio decreased progressively during transition from the vegetative to the reproductive state of growth, from a = 1.48 initially to a = 1.15 six days after the beginning of induction. The divergence angle was not altered during the transition. This change in phyllotaxis is interpreted as a change in the relative positioning of leaf primordia on the transitional apex. This transient change appears to be identical with the previously described long-term change of the phyllotaxis of Xanthium brought about by treatment of plants with gibberellic acid.     

Coastal eutrophication: recent developments in definitions and implications for monitoring strategies

The word ‘eutrophication’ has its root in two Greekwords: ‘eu’ which means ‘well’ and ‘trope’which means ‘nourishment’. The modern use of theword eutrophication is related to inputs and effects of nutrientsin aquatic systems. Despite a common understanding of its causesand effects, there is no agreed definition of coastal eutrophication.This communication aims to review recent developments in thedefinitions of coastal eutrophication, all of which focus on‘accelerated growth’, and to discuss the implicationsin relation to monitoring and assessment of ecological status.It is recommended that measurements of primary production, beinga sensitive and accurate indicator of eutrophication, shouldbe mandatory when monitoring and assessing the ecological statusof coastal waters. Written responses to this article should be submitted to KevinFlynn at k.j.flynn@swansea.ac.uk within two months of publication.For further information, please see the Editorial ‘Horizons’in Journal of Plankton Research, Volume 26, Number 3, Page 257.     

The Growth and Carbon Economy of Selection Lines of Lolium perenne cv. S23 with Differing Rates of Dark Respiration: 1. Grown as Simulated Swards During a Regrowth Period

Simulated swards of each of two selection lines of Lolium perennecv. S23 with ‘fast’ and ‘slow’ ratesof ‘mature tissue’ respiration were establishedin growth rooms at 20/15 °C day/night temperatures and studiedover four successive regrowth periods of 46, 30, 26 and 53 daysduration. The ‘slow’ line outyielded the ‘fast’,both in harvestable shoot (above a 5 cm cut) and in root andstubble. Its advantage increased over successive regrowth periodsto 23 per cent (total biomass). Gas analysis measurements onthe entire communities (including roots), during the final regrowthperiod, showed that the ‘slow’ line had a 22–34per cent lower rate of dark respiration per unit dry weight.This enabled it to maintain its greater mass of tissue for thesame cost in terms of CO₂ efflux per unit ground area. Halfthe extra dry weight produced by the ‘slow’ line,relative to the ‘fast’, could be attributed to itsmore economic use of carbon. The rest could be traced to a 25per cent greater tiller number which enabled the ‘slow’line to expand leaf area faster (though not at a greater rateper tiller), intercept more light and fix more carbon, earlyin the regrowth period. Lolium perenne L., ryegrass, respiration, maintenance respiration, tiller production, simulated swards, canopy photosynthesis, carbon economy     

Plant Morphology: The Historic Concepts of Wilhelm Troll, Walter Zimmermann and Agnes Arber

Recent molecular systematic and developmental genetic findingshave drawn attention to plant morphology as a discipline dealingwith the phenotypic appearance of plant forms. However, sincedifferent terms and conceptual frameworks have evolved overa period of more than 200 years, it is reasonable to surveythe history of plant morphology; this is the first of two paperswith this aim. The present paper deals with the historic conceptsof Troll, Zimmermann and Arber, which are based on Goethe'smorphology. Included are contrasting views of ‘unity anddiversity’, ‘position and process’, and ‘morphologyand phylogeny’, which, in part, are basic views of currentplant morphology, phylogenetic systematics and developmentalgenetics. Wilhelm Troll established the ‘type concept’and the ‘principle of variable proportions’. Hehas provided the most comprehensive overview of the positionalrelations of plant forms. Agnes Arber started from the universaldynamics of life and attempted to describe all structures asprocesses. She paid attention to ‘repetitive branching’,‘differential growth’, and ‘parallelism’.As a result she has recently been rediscovered by developmentalbotanists. Walter Zimmermann rejected any metaphysical influenceon plant form and instead called for objective procedures. Hewas mainly interested in phylogenetic ‘character transformation’and the ‘reconstruction of genealogical lines’.Guided by the example of flower-like inflorescences, a futurepaper will deal with functional and developmental constraintsinfluencing plant forms. Recent morphological concepts (‘trialectical’,‘continuum’/‘fuzzy’, ‘processmorphology’) will be discussed and related to currentmorphological and developmental genetic research. Copyright2001 Annals of Botany Company Plant form, plant morphology, natural philosophy, homology, phylogeny, Goethe, Troll, Arber, Zimmermann, typology, character transformation, differential growth, complementarity     

Properties of maximal spacing on a circle related to phyllotaxis and to the golden mean

A class of divergence angles φ_G of phyllotaxis is defined that distribute leaves about the stem of a plant in a more uniform manner than do nearby angles. A theorem of Swierczkowski concerning the intervals between adjacent points placed on a circle according to the divergence angle 137·5° is generalized to include the other angles encountered in phyllotaxis. These angles are characterized by having continued fraction expansions containing no intermediate fractions after a finite number of terms. This criterion is shown to be sufficient for uniform spacing of leaves.A morphogen concentration field established by the leaves acting as sources is determined principally by :he geometrical spacing of the leaves and hence by their divergence angle. It is shown that the mean square of such a concentration field, is a relative minimum if the leaves are positioned by means of one of the φ_G. Thornley's dynamic scheme for determining the phyllotaxis divergence angles by positioning a new leaf at the minimum of the concentration field of previously placed leaves is also shown to be related to the spacing properties of the φ_G.     

The Polarity of the Cell-wall of Valonia

Further evidence is presented to show that the systems of orientationof cellulose fibrils in the cell-walls of Valonia and Dictyosphaeriamay be described by reference to two ‘poles’ uponwhich these orientation systems converge. The ‘poles’are real structural features of the wall.     

The Growth and Carbon Economy of Selection Lines of Lolium perenne cv. S23 with Differing Rates of Dark Respiration: 2. Grown as Young Plants from Seed

Young plants of two selection lines of Lolium perenne cv. S23with ‘fast’ and ‘slow’ rates of ‘maturetissue’ respiration were individually grown from seed,together with plants of S23, their common parent, in 9.2 cmpots in a controlled environment at 20/15 °C day/night temperatures. No significant differences were found between the genotypesin leaf extension and tiller production during this early stageof their growth. They did differ however, by an average of 26%,in the rate of dark respiration of fully expanded leaf laminae.The use of a simple model demonstrated that such a differencein respiration could alone account for the different rates ofdry matter production shown by the selection lines when grownas young crops from seed. Possible penalties of ‘slow’respiration are also considered. Lolium perenne L., ryegrass, respiration, maintenance respiration, stimulated swards, leaf growth, tiller production, carbon economy     

CAPITULUM PHYLLOTAXIS AND NUMERICAL CANALIZATION IN MICROSERIS PYGMAEA (ASTERACEAE: LACTUCEAE)

The positions at which floret primordia arise in developing capitulum buds of Microseris pygmaea D. Don have been mapped by computer-assisted light microscopy. The primordia can be assigned positions along a basic phyllotactic spiral with a divergence angle of about 137.5°. In addition, there are regular deviations from a spiral arrangement. Typically, the first 26 primordia in phyllotactic sequence are arranged in two concentric circles of 13 primordia with considerable deviations in the divergence angle and in the distances between primordia along a parastichy at positions 13 and 26. This arrangement can be simulated by geometric models that include nearest neighbor packing, together with spiral phyllotaxis. The circular arrangement of peripheral primordia at nearly equal radial distances from the center of the developing capitulum helps to explain the numerical constancy (canalization) of peripheral structures, especially the constant number of 13 inner phyllaries on heads with very different numbers of florets.     

An interpretation of Fujita's frequency diagrams in phyllotaxis

Fujita's diagrams in phyllotaxis, showing the frequencies of divergence angles as a function of these angles for low phyllotactic patterns such as (2, 1) and (3, 2), which are approximately normal curves centered at the limitdivergence angle of 137.51°, are shown to be puzzling when compared to results and observations in the field. An analysis of these diagrams is proposed, in the context of Fujita's methodology, of data from other sources, of a mathematical theorem on lattices, and of the contact pressure theory of phyllotaxis.     

Diurnal Phototropism in Leaves of Lavatera cretica L. under Conditions of Simulated Solar-Tracking

Diurnal laminar reorientation was followed in solar-trackingleaves of Lavatera cretica L. under simulated conditions. Asimulated ‘sun’ was moved over the lamina in a 180?arc in the vertical plane of the mid-vein, at an angular velocityof 15? h^–1 in a regime of 12-h photoperiods. In one groupof plants the petioles of the experimental leaves were arrangedto face ‘sunrise’, while in the other they werearranged to face ‘sunset’. At ‘sunrise’,the laminae in both groups, which were inclined towards theanticipated direction of ‘sunrise’, changed theirelevation towards the rising ‘sun’, resulting inprogressive reduction in the angle of incidence (AI) of lighton the laminar surface (AI= differential between laminar and‘solar’ elevation). As a result, laminar and ‘solar’elevation converged, and laminar reorientation gradually ceased,until the ‘solar’ elevation had passed the normalto the laminar surface (AI=0?). laminar reorientation was thenresumed, but its direction was reversed to follow the directionof ‘solar’ reorientation. During most of the remaining‘day’, laminar elevation (LE) trailed that of the‘sun’ by an average of 11?-14?. Laminar reorientationthen anticipated ‘sunset’ by starting to slow down60 to 90min in advance. During the 12-h dark period, the laminareoriented towards the anticipated direction of the subsequent‘sunrise’. The time-course of nocturnal reorientationwas qualitatively different in the two groups of experimentalplants. The time-course of diurnal phototropism under naturaland simulated conditions is analysed and compared and differencesand similarities between them are discussed. Key words: Diurnal phototropism, solar-tracking, vectorial excitation     

Effects on Growth and Development of Individual Chromosomes from Slow-growing Lophopyrum elongatum Love when Incorporated into Bread Wheat (Triticum aestivum L.)

Aspects of growth and development were evaluated in the fast-developingannual Triticum aestivum L. ‘Chinese Spring’, theslow-developing perennial Lophopyrum elongatum Löve, theiramphiploid, and chromosome addition and substitution lines ofL. elongatum into ‘Chinese Spring’. Relative growthrates (RGR) of shoots of L. elongatum and the amphiploid werelower than those of ‘Chinese Spring’ (34 and 13%respectively) and main stem development was also slower. Therewas no difference in shoot RGR of any of the chromosome additionor substitution lines and that of ‘Chinese Spring’when assessed between Haun stages 2.0 and 5.0. In contrast,several aspects of plant development were observed to differin the chromosome addition and substitution lines. SubstitutingE genome chromosomes (with the exceptions of 3E and 4E) forD genome chromosomes, or adding E genome chromosomes, slowedthe rate of main stem development, at least up to Haun stage5.0. Despite these differences in the rate of main stem development,the appearance of adventitious roots commenced at approximatelyHaun stage 2.0 in all genotypes. However, the numbers of adventitiousroots and tillers at the 5.0 Haun stage differed between someof the lines when compared to ‘Chinese Spring’.Although incorporation of some L. elongatum chromosomes alteredaspects of plant development, all lines showed more similarityto bread wheat than to L. elongatum, reflecting, in part, thegreater genetic contribution made by bread wheat to these lines.Copyright 2001 Annals of Botany Company Adventitious roots, chromosome addition and substitution lines, Haun stage, Lophopyrum elongatum, relative growth rate (RGR), Triticum aestivum(wheat)