Effects of fire frequency on oak litter decomposition and nitrogen dynamics

Rapid speciation within some plant families has been attributed to the evolution of floral spurs and to the effect of spur length on plant reproductive success. The flowers of Impatiens capensis (jewelweed) possess a long, curved spur in which nectar is produced and stored. Spur length and curvature varies among plants within one population. Here I document that spur shape is variable in natural populations, variation within plants is less than variation among plants, and spur shape is correlated with components of female and male reproductive success. The apparent natural selection is weakly directional in 1 of 2 years, with greatest seed production and pollen removal occurring in flowers with the greatest spur curvature. Bee pollinator visit length is longest at flowers with highly curved spurs, and they leave less nectar in these spurs than in flowers with straighter spurs. Spur angle evolution may be limited, at least in part, by opposing selection by nectar-robbers who prefer to visit flowers with greater spur curvature. Other factors that might contribute to the maintenance of spur angle variation are temporal variation in the strength of selection and potential genetic correlations of spur shape with other traits under selection.     

Increases in leaf nitrogen concentration and leaf area did not enhance spur survival and return bloom in almonds (<Emphasis Type="Italic">Prunus dulcis</Emphasis> [Mill.] DA Webb)

Mature almond trees bear fruit mainly on short shoots called spurs, with only a small percentage of fruit produced laterally on long 1-year-old shoots. As a result, maintenance of large numbers of healthy spurs per tree is critical for fruit production. However, spurs that bear fruit have lower leaf area, leaf nitrogen content, and CO₂ assimilation rate than non-fruiting spurs. This has been correlated with reduced percentages of spur survival and return bloom the following season. Thus, we hypothesized that spur leaf area, and ultimately spur health could be enhanced through application of foliar sprays and soil nitrogen treatments that would enhance leaf nitrogen content and spur leaf area. To test our hypothesis, we selected almond trees exhibiting significant yield differences as a consequence of differential soil rates of nitrogen fertilization (N rate) for three prior years (140, 224, and 392 kg/ha). In each tree, three spur types [non-fruiting spurs (F0); spurs with one fruit (F1); spurs with two fruit (F2)] were selected on the east side of the canopy and tracked for one complete season (2011–2012). Four foliar treatments (nutrient replacement, nutrient replacement with biostimulant, nitrogen, and non-spray) were directly applied to individual spurs in each N rate in the spring of 2011 and characteristic such as leaf nitrogen, and fruit quality were recorded throughout the season. In winter of 2012, spur survival and return bloom were addressed through individual visual inspection of the tagged spurs. In this experiment, soil and foliar N treatments effectively increased spur leaf area, fruit, and leaf nitrogen concentration. In the high N treatment, the leaf nitrogen values exceeded the critical nitrogen concentration established for almond trees and the critical leaf area for spur survival and blooming thresholds established by past research in this area. However, none of these positive changes in leaf N or leaf area improved spur survival and/or return boom of any spur type. Indeed, survival and hull + shell weight tended to be lower in the population of fruiting spurs with the highest leaf area and leaf nitrogen concentration and the return bloom probabilities were always lower in fruiting spurs than in non-fruiting spurs, independent of the nitrogen rate. These results and the relationship between nitrogen rate and spur survival are discussed.     

LONG-TONGUED FLY POLLINATION AND EVOLUTION OF FLORAL SPUR LENGTH IN THE DISA DRACONIS COMPLEX (ORCHIDACEAE)

Field studies in South Africa showed that floral spur length in the Disa draconis complex (Orchidaceae) varies enormously between populations in the southern mountains (means = 32–38 mm), lowland sandplain (mean = 48 mm), and northern mountains (means = 57–72 mm). We tested the hypothesis that divergence in spur length has resulted from selection exerted through pollinator proboscis length. Short-spurred plants in several southern mountain populations, as well as long-spurred plants in one northern mountain population, were pollinated by a horsefly, Philoliche rostrata (Tabanidae), with a proboscis length that varied from 22 to 35 mm among sites. Long-spurred plants on the sandplain were pollinated by the tanglewing fly, Moegistorynchus longirostris (Nemestrinidae), which has a very long proboscis (mean = 57 mm). Selection apparently favors long spurs in sandplain plants, as artificial shortening of spurs resulted in a significant decline in pollen receipt and fruit set, although pollinaria removal was not significantly affected. Fruit set in the study populations was limited by pollen availability, which further suggests that selection on spur length occurs mainly through the female component of reproductive success.     

Pollinators and nectar robbers cause directional selection for large spur circle in Impatiens oxyanthera (Balsaminaceae)

Nectar spurs have an important role in floral evolution and plant–pollinator coadaptation. The flowers of some species possess spurs curving into a circle. However, it is unclear whether spur circle diameter is under direct selection pressure from different sources, such as pollinators and nectar robbers. In this study, we quantified selection on some floral traits, such as spur circle diameter in Impatiens oxyanthera (Balsaminaceae) using phenotypic selection analysis and compared the relative importance of pollinators and nectar robbers as selective agents using mediation analysis. The study showed that pollinators caused significant selection on corolla length, spur curvature and spur circle diameter while nectar robbers only imposed strong selection on spur circle diameter. Pollinators favored flowers with large corolla, curly spurs and large spur circle while nectar robbers preferred flowers with small spur circle. More pollinator visits resulted in higher female reproductive success, while robbery reduced female fitness. Conflicting selection on spur traits from pollinators and nectar robbers was not found. Mediation analysis showed that selection on floral traits through nectar robbing was stronger than selection through pollination. The results suggested that pollinators and nectar robbers jointly mediated the directional selection for large spur circle, and nectar robbers caused stronger selection than pollinators on floral traits.     

ROOT GLOCHIDS AND ROOT SPURS OF OPUNTIA ARENARIA (CACTACEAE)

Root glochids of Opuntia arenaria Engelm. are produced by adventitious buds (areoles) that arise endogenously on fleshy roots. If they remain active, root areoles become dwarf shoots that initiate only glochids and trichomes unless the roots are uncovered or the shoots bearing the roots are removed. Then they will expand, producing aerial shoot joints. Root spurs are formed in great numbers on the profusely branched system of small roots. They consist of clusters of rootlets about one mm long that are initiated in sympodial sequence, each new rootlet arising endogenously near the base of the preceding one. On older spurs this results in the formation of a short axial peg. Rootlet primordia lack a root cap from the beginning and appear to mature quickly, losing all their meristematic characteristics and becoming completely covered with root hairs, even over the tip. They may represent a mechanism for the rapid production of root hairs during the infrequent periods when moisture is available. In any case, they seem to be transitory structures because cork soon forms beneath the older ones on a spur.     

Evolution of spur-length diversity in Aquilegia petals is achieved solely through cell-shape anisotropy

The role of petal spurs and specialized pollinator interactions has been studied since Darwin. Aquilegia petal spurs exhibit striking size and shape diversity, correlated with specialized pollinators ranging from bees to hawkmoths in a textbook example of adaptive radiation. Despite the evolutionary significance of spur length, remarkably little is known about Aquilegia spur morphogenesis and its evolution. Using experimental measurements, both at tissue and cellular levels, combined with numerical modelling, we have investigated the relative roles of cell divisions and cell shape in determining the morphology of the Aquilegia petal spur. Contrary to decades-old hypotheses implicating a discrete meristematic zone as the driver of spur growth, we find that Aquilegia petal spurs develop via anisotropic cell expansion. Furthermore, changes in cell anisotropy account for 99 per cent of the spur-length variation in the genus, suggesting that the true evolutionary innovation underlying the rapid radiation of Aquilegia was the mechanism of tuning cell shape.     

The relationship between cell division and elongation during development of the nectar-yielding petal spur in Centranthus ruber (Valerianaceae)

Background and Aims Floral spurs are hollow, tubular outgrowths that typically conceal nectar. By their involvement in specialized pollinator interactions, spurs have ecological and evolutionary significance, often leading to speciation. Despite their importance and diversity in shape and size among angiosperm taxa, detailed investigations of the mechanism of spur development have been conducted only recently.Methods Initiation and growth of the nectar-yielding petal spur of Centranthus ruber ‘Snowcloud’ was investigated throughout seven stages, based on bud size and developmental events. The determination of the frequency of cell division, quantified for the first time in spurs, was conducted by confocal microscopy following 4'',6-diamidino-2-phenylindole (DAPI) staining of mitotic figures. Moreover, using scanning electron microscospy of the outer petal spur surface unobstructed by trichomes, morphometry of epidermal cells was determined throughout development in order to understand the ontogeny of this elongate, hollow tube.Key Results Spur growth from the corolla base initially included diffuse cell divisions identified among epidermal cells as the spur progressed through its early stages. However, cell divisions clearly diminished before a petal spur attained 30 % of its final length of 4·5 mm. Thereafter until anthesis, elongation of individual cells was primarily responsible for the spur’s own extension. Consequently, a prolonged period of anisotropy, wherein epidermal cells elongated almost uniformly in all regions along the petal spur’s longitudinal axis, contributed principally to the spur’s mature length.Conclusions This research demonstrates that anisotropic growth of epidermal cells – in the same orientation as spur elongation – chiefly explains petal spur extension in C. ruber. Representing the inaugural investigation of the cellular basis for spur ontogeny within the Euasterids II clade, this study complements the patterns in Aquilegia species (order Ranunculales, Eudicots) and Linaria vulgaris (order Lamiales, Euasterids I), thereby suggesting the existence of a common underlying mechanism for petal spur ontogeny in disparate dicot lineages.     

Modelling three-dimensional flow over spur-and-groove morphology

Spur-and-groove (SAG) morphology characterizes the fore reef of many coral reefs worldwide. Although the existence and geometrical properties of SAG have been well documented, an understanding of the hydrodynamics over them is limited. Here, the three-dimensional flow patterns over SAG formations, and a sensitivity of those patterns to waves, currents, and SAG geometry were characterized using the physics-based Delft3D-FLOW and SWAN models. Shore-normal shoaling waves over SAG formations were shown to drive two circulation cells: a cell on the lower fore reef with offshore flow over the spurs and onshore flow over the grooves, except near the seabed where velocities were always onshore, and a cell on the upper fore reef with offshore surface velocities and onshore bottom currents, which result in depth-averaged onshore and offshore flow over the spurs and grooves, respectively. The mechanism driving this flow results from the net of the radiation stress gradients and pressure gradient, which is balanced by the Reynolds stress gradients and bottom friction that differ over the spur and over the groove. Waves were the primary driver of variations in modelled flow over SAG, with the flow strength increasing for increasing wave heights and periods. Spur height, SAG wavelength, and the water depth at peak spur height were the dominant influences on the hydrodynamics, with spur heights directly proportional to the strength of SAG circulation cells. SAG formations with shorter SAG wavelengths only presented one circulation cell on the shallower portion of the reef, as opposed to the two circulation cells for longer SAG wavelengths. SAG formations with peak spur heights occurring in shallower water had stronger circulation than those with peak spur heights occurring in deeper water. These hydrodynamic patterns also likely affect coral and reef development through sediment and nutrient fluxes.

     

Spur development in chick embryos

Normal development of spurs and horny squamae have been studied in histological preparations obtained from the skin of tarsometatarsus in 8--16-day-old embryos. During the first, initial stage of development, by means of rearrangement of cellular matter and cell migration, three main parts of the spur are layed down -- the horny cover, the spur body and the fibrovascular cushion. For the second stage, vigorous growth of the spur germ at the expense of proliferative activity of its cells is characteristic. At the third stage (after hatching) in males, the spur body outgrows and the bony core is formed. Morphogenesis of the horny squama can be devided into two stages. At the initial stages by means of condensing cellular elements, squamous papilla and horny shell are layed down. The fibrovascular cushion is absent. The second stage is similar to the spur one and is characterized by growth of all the germ parts at the expense of cell proliferation. Comparing morphogeneses of the squama and the spur, it is possible to conclude that phylogenetic transformation of the squama into the spur is performed by two means (modi) of phyloembryogenesis: by means of adding new signs of development to the initial terminal stages of its morphogenesis.     

Manipulation of trait expression and pollination regime reveals the adaptive significance of spur length

Understanding the mechanisms of adaptive population differentiation requires that both the functional and adaptive significance of divergent traits are characterized in contrasting environments. Here, we (a) determined the effects of floral spur length on pollen removal and receipt using plants with artificial spurs representing the species-wide variation in length, and (b) quantified pollinator-mediated selection on spur length and three traits contributing to floral display in two populations each of the short-spurred and the long-spurred ecotype of the orchid Platanthera bifolia. Both pollen receipt and removal reached a maximum at 28–29 mm long spurs in a short-spurred population visited by short-tongued moths. In contrast, pollen receipt increased linearly across the tested range (4–52 mm) and pollen removal was unrelated to spur length in a long-spurred population predominantly visited by a long-tongued moth. The experimentally documented effects on pollen transfer were not reflected in pollinator-mediated selection through female fitness or pollen removal indicating that the natural within-population variation in spur length was insufficient to result in detectable variation in pollen limitation. Our study illustrates how combining trait manipulation with analysis of causes and strength of phenotypic selection can illuminate the functional and adaptive significance of trait expression when trait variation is limited.     

Hawkmoth pollination of aerangoid orchids in Kenya, with special reference to nectar sugar concentration gradients in the floral spurs

The African orchid flora has a high proportion of species with long-spurred white flowers. Few data exist to test the prediction that this floral syndrome pattern reflects an important role for hawkmoth pollination in the evolution and ecology of these orchids. The pollination biology of five aerangoid orchid species (Rangaeris amaniensis, Aerangis brachycarpa, A. confusa, A. thomsonii, and A. kotschyana) was investigated in Kenya. Four of these have long spurs (>10 cm) and were pollinated by Agrius convolvuli and Coelonia fulvinotata. Aerangis confusa, which has relatively short spurs (ca. 4 cm), was pollinated by the short-tongued hawkmoths Hippotion celerio and Daphnis nerii. Nectar frequently filled the entire spur in some of the study species, even at anthesis. Sugar concentration of the nectar of four species was found to vary from ca. 1% at the mouth of the spur to 20% at the tip. Gradients were expressed more strongly in species with long, straight spurs. Species with spirally twisted spurs showed both steep and shallow nectar gradients. These gradients, previously unknown in plants, may function as a "sugar trail," enticing long-tongued hawkmoths to probe deeply into spurs without incurring the cost of filling an entire spur with concentrated nectar. In addition, the most concentrated nectar is kept out of reach of short-tongued pollinators.     

Comparison of stochastic and deterministic methods for modeling spur kinetics

Stochastic and deterministic kinetic methods have been used to model the temporal evolution of spatially nonhomogeneous clusters of reactants resulting from the dissociation of one to six water molecules into either H3O+, OH, and e-aq, or H atoms and OH radicals. When the ionic fragmentation initially producing H3O+, OH, and e-aq is considered, the stochastic and deterministic methods predict similar time dependences for the decay of the reactive species; however, the two methods suggest very different product yields. For a two-dissociation spur, the deterministic treatment overestimates both the H2 and the H2O2 yields by about 70%. The error decreases to less than 15% for a spur with six water dissociations. For a distribution of spurs representing a high-energy electron track, the differences in the predicted yields of reactants are less than 6% at 0.1 microseconds, but the stochastic and deterministic predictions for the yields of H2 and H2O2 differ by about 50%. The kinetics of spurs produced by the fragmentation of water to H atom and OH radical shows discrepancies in both the reactant and the product yields. The size of the discrepancy decreases as the number of H/OH pairs increases, and the predictions of the two techniques are almost the same for clusters of six water dissociations.     

Pollinators of the Rocky Mountain columbine: temporal variation, functional groups and associations with floral traits

Background and Aims

Pollinators together with other biotic and some abiotic factors can select for floral traits. However, variation in pollinator abundance over time and space can weaken such selection. In the present study, the variation in pollinator abundance over time and space was examined in populations of the Rocky Mountain columbine. The variation in three floral traits is described and correlations between pollinator type, functional pollinator groups or altitude and floral traits are examined.

Methods

Pollinator observations took place in six Aquilegia coerulea populations over 1–4 years and spur length, flower colour and sepal length were measured in 12 populations. Pollinator abundance, measured as visits per flower per hour, was compared among populations and years. Pollinators were grouped into two functional groups: pollen or nectar collectors. The following associations were examined: annual presence of hawkmoths and whiter flowers with longer spurs; the presence of Sphinx vashti and longer spurs; and higher altitudes and whiter flowers. The study looked at whether an increase in the proportion of hawkmoths in a population was associated with whiter and larger flowers with longer spurs.

Key Results

The abundance of different pollinator groups varied over time and space. Floral traits varied among populations. Higher altitude was correlated with bluer flowers. Whiter flowers were associated with the annual presence of hawkmoths. Populations visited by Sphinx vashti had longer spurs than populations visited only by Hyles lineata. Populations with greater percentage of nectar-collecting pollinators did not have whiter, larger flowers with longer spurs.

Conclusions

Despite the large variation in pollinator abundance over time and space, one species of bumble-bee or hawkmoth tended to predominate in each population each year. Future studies of Aquilegia coerulea should examine the specific influences of pollinators and the environment on flower colour and of hawkmoth species on spur length.Key words: Aquilegia coerulea, columbine, pollinator abundance, bumble-bee, hawkmoth, flower colour, spur length, functional pollinator group, altitude, floral trait     

Comparative Studies on Physiological and Morphological Features of Bearing and Non-Bearing Spurs of the Apple Tree II. The Effect of Fruiting and Growth Regulator Sprays on Respiration Mate of Leaves

The respiration rate was investigated in leaves of bearing and non-bearing spurs of biennially fruiting apple trees, Landsberger Reinette cultivar, with a Warburg apparatus. Trees in “off year” were sprayed with gibberellic acid (GA) and those in “on year” with succinic acid-2,2-dimethylhydra-zide (SADH) and naphthaleneacetic acid (NAA). Two-year results showed that the respiration rate of leaves of bearing spurs was 10 to 25 per cent higher than that of the non-bearing spur leaves. GA increased the respiration of leaves of non-bearing spurs about 25 per cent, on an average. SADH and NAA greatly decreased the respiration of bearing spur leaves; the latter about 30 per cent of that of the controls. At the same time SADH and NAA caused a marked reduction of growth in terminal shoots of trees in “on year” while GA increased the growth in “off year” trees about 50 per cent. The surface of the spur leaves was inversely correlated with the respiration rates. It is suggested that the low respiration rate of spur leaves may be a condition for flower bud formation.     

Are spurred cyathia a key innovation? Molecular systematics and trait evolution in the slipper spurges (Pedilanthus clade: Euphorbia, Euphorbiaceae)

The study of traits that play a key role in promoting diversification is central to evolutionary biology. Floral nectar spurs are among the few plant traits that correlate with an enhanced rate of diversification, supporting the claim that they are key innovations. Slight changes in spur morphology could confer some degree of premating isolation, explaining why clades with spurs tend to include more species than their spurless close relatives. We explored whether the cyathial nectar spur of the Pedilanthus clade (Euphorbia) may also function as a key innovation. We estimated the phylogeny of the Pedilanthus clade using one plastid (matK) and three nuclear regions (ITS and two G3pdh loci) and used our results and a Yule model of diversification to test the hypothesis that the cyathial spur correlates with an increased diversification rate. We found a lack of statistical support for the key innovation hypothesis unless specific assumptions regarding the phylogeny apply. However, the young age (hence small size) of the group may limit our ability to detect a significant increase in diversification rate. Additionally, our results confirm previous species designations, indicate higher homoplasy in cyathial than in vegetative features, and suggest a possible Central American origin of the group.     

Critical periods in the development of spurs in light of the theory of phylembryogenesis]

Two critical periods have been established in the development of spurs representing a derivative of the corneous squama. During the first period the exposure to x-rays results in the development of the corneous squama instead of the spur, and the same exposure during the second critical period completely suppresses the development of the spur. These periods seem to correspond to the truning-points in the chain of morphogenetical processes of the spur formation associated with phylembryogeneses. Department of Embryology, Institute of Experimental Medicine, USSR Acad. Med. Sci., Leningrad.     

Avian spurs

Spur size and number in the various species of Galliformes are considered in relation to body size and social system. Spur length is absolutely and relatively greater among large tropical species. Multiple-spurred species are intermediate in size between spurless and single-spurred species, but have longer legs. Possession of spurs is common, and presumably originated among monogamous species, but is still commoner among polygamous ones. Different spur shapes possibly reflect wounding versus stunning strategies of fighting. Wing spurs in other birds are briefly compared.     

Mating system correlates of tarsal spurs in the Phasianidae

Two methods for the analysis of cross-species data are used to investigate whether tarsal spur length or number are correlated with mating system in three phasianid subfamilies. One method, the phylogenetic regression. suggests that polygamous species have relatively longer and fewer spurs than monogamous species. hut this relationship is not found when a matched-pairwise comparison is used. These ambiguous results suggest that at the moment it is not possible to conclude either way whether spurs are under any selective pressure related to mating system, and that previously published conclusions are not supported.