Difference in cuticular transpiration and sclerophylly in juvenile and adult pine needles relates to the species-specific rates of development

Pinus species show remarkable ontogenetic differences in needle morphology (heterophylly) between juvenile and adult vegetative phases. This developmental shift may play an adaptative role in their success under diverse habitats. As a first step to know the functional differences between each vegetative phase, we compared water loss through the cuticles of juvenile and adult needles of 21-month-old nursery-grown seedlings of nine hard pine species. Cuticular transpiration (CT), calculated after complete stomatal closure, was obtained by leaf-drying curves, and was related to leaf, ontogenetic and climatic parameters. The rate of cuticular transpiration (RCT) between juvenile and adult needles differed across pine species, and in particular segregated the Mediterranean species Pinus canariensis and P. halepensis, from the Eurasian P. uncinata and introduced species P. radiata. For these species, RCT was always higher in juvenile needles. The different leaf and ontogenic parameters studied were correlated with the variation in RCT among the nine pine species. We discuss this relationship in the light of the species ecology. Besides their possible adaptive interpretation, these results suggest an underlying need to consider the ontogenetic heterophylly when assessing functional traits in hard pine seedlings, in particular those traits that govern water relations.     

Complex population genetic structure in the endemic Canary Island pine revealed using chloroplast microsatellite markers

The Canary archipelago, located on the northwestern Atlantic coast of Africa, is comprised of seven islands aligned from east to west, plus seven minor islets. All the islands were formed by volcanic eruptions and their geological history is well documented providing a historical framework to study colonization events. The Canary Island pine (Pinus canariensis C. Sm.), nowadays restricted to the westernmost Canary Islands (Gran Canaria, Tenerife, La Gomera, La Palma and El Hierro), is considered an old (Lower Cretaceous) relic from an ancient Mediterranean evolutionary centre. Twenty seven chloroplast haplotypes were found in Canary Island pine but only one of them was common to all populations. The distribution of haplotypic variation in P. canariensis suggested the colonization of western Canary Islands from a single continental source located close to the Mediterranean Basin. Present-day populations of Canary Island pine retain levels of genetic diversity equivalent to those found in Mediterranean continental pine species, Pinus pinaster and Pinus halepensis. A hierarchical analysis of variance (AMOVA) showed high differentiation among populations within islands (approximately 19%) but no differentiation among islands. Simple differentiation models such as isolation by distance or stepping-stone colonization from older to younger islands were rejected based on product-moment correlations between pairwise genetic distances and both geographic distances and population-age divergences. However, the distribution of cpSSR diversity within the islands of Tenerife and Gran Canaria pointed towards the importance of the role played by regional Pliocene and Quaternary volcanic activity and long-distance gene flow in shaping the population genetic structure of the Canary Island pine. Therefore, conservation strategies at the population level are strongly recommended for this species.Communicated by D.B. NealeA. Gómez and S.C. González-Martínez as joint authors     

Chloroplast microsatellites reveal colonization and metapopulation dynamics in the Canary Island pine

Chloroplast microsatellites are becoming increasingly popular markers for population genetic studies in plants, but there has been little focus on their potential for demographic inference. In this work the utility of chloroplast microsatellites for the study of population expansions was explored. First, we investigated the power of mismatch distribution analysis and the F(S) test with coalescent simulations of different demographic scenarios. We then applied these methods to empirical data obtained for the Canary Island pine (Pinus canariensis). The results of the simulations showed that chloroplast microsatellites are sensitive to sudden population growth. The power of the F(S) test and accuracy of demographic parameter estimates, such as the time of expansion, were reduced proportionally to the level of homoplasy within the data. The analysis of Canary Island pine chloroplast microsatellite data indicated population expansions for almost all sample localities. Demographic expansions at the island level can be explained by the colonization of the archipelago by the pine, while population expansions of different ages in different localities within an island could be the result of local extinctions and recolonization dynamics. Comparable mitochondrial DNA sequence data from a parasite of P. canariensis, the weevil Brachyderes rugatus, supports this scenario, suggesting a key role for volcanism in the evolution of pine forest communities in the Canary Islands.     

THE EFFECT OF SUGARS AND APPLIED CHEMICALS ON HETEROBLASTIC DEVELOPMENT IN IPOMOEA PURPUREA GROWN IN ASEPTIC CULTURE

Seedlings of Ipomoea purpurea were grown in aseptic culture in order to investigate more directly than had been possible in pot culture the effect of carbohydrate and protein nutrition on heteroblastic development which in this species involves a change from entire juvenile to lobed adult leaves. The effect of sugars was to accelerate the development of the adult leaf form while casein hydrolysate retarded it. The growth retardants, AMO-1618 and CCC, also promoted the formation of the adult leaf. These results are discussed in the light of current views on the factors controlling heteroblastic development.     

Age and seasonal effects on predator-sensitive foraging in squirrel monkeys (Saimiri sciureus): a field experiment

A field experiment was conducted to examine the effect of perceived predation risk on the use of foraging areas by juvenile and adult primates under different conditions of local food abundance. Wild squirrel monkeys, Saimiri sciureus, were observed in an experiment conducted during the dry and the wet seasons at a site in Eastern Amazonia, Brazil. Animals were presented with feeding platforms that differed in food quantity and exposure to aerial predators through varying vegetative cover. In the dry season, juveniles and adults chose platforms based solely on food quantity. However, in the wet season, juveniles foraged preferentially on high-reward platforms only if cover level also was high (i.e., potentially offered greater concealment from predators). In contrast, adults showed the same pattern of platform use regardless of season. These results indicate that age and local resource availability based on seasonality affect whether primates forage in a predator-sensitive manner. Juveniles may be more sensitive to predation risk when foraging, and individuals may take fewer risks when resource abundance is high in their environment.     

Plant structural defences against browsing birds: a legacy of New Zealand's extinct moas

Browsing by large vertebrates has been a major force in the evolution of terrestrial plants but Holocene extinctions of the browsers have left a legacy of broken biotic partnerships. Ratite birds were the largest herbivores in several regions, such as the moas of New Zealand. Many woody plants there have a distinct form of branching, described as "divaricate", with thin, wide angled, branches intertwining to form a tangled canopy. Divaricate branching has been interpreted as a form of protection against climate extremes or as an anachronistic defense against the extinct moas. Here we report the first experimental evidence that many of these plants are defended against extant ratite browsers. In feeding experiments on two tree species with different (heteroblastic) juvenile and adult branch morphology, emus and ostriches obtained adequate feeding rates from adult shoots but sub-maintenance feeding rates from juvenile shoots with the ratite-resistant traits. Divaricate juvenile shoots suffered 30–70% less biomass removal to the birds than adult shoots. Ratites browse by a distinctive clamping and tugging action. Structural defence traits that exploit the limitations of this feeding mode include narrow, strong, elastic branches that resist being torn off, wide branching angle ("divaricate") that makes shoots difficult to swallow, and small, widely spaced leaves. This novel plant architectural defence has developed in at least 20% of the native woody flora of New Zealand, including 10 heteroblastic tree species that exhibit the ratite-resistant strategy until they reach ca 3 metres height. It is also a major axis of variation amongst homoblastic woody shrub species. The defences are useless against mammalian browsers that shear shoots, contributing to marked decreases in the abundances of ratite-resistant species in New Zealand after the introduction of mammals.     

Leaf heteroblasty is not an adaptation to shade: seedling anatomical and physiological responses to light

Heteroblastic plants produce markedly different leaf morphologies between juvenile and adult stages, while homoblastic plants exhibit little or gradual changes. We tested the hypothesis that the leaf morphology of the seedling stage of New Zealand heteroblastic species is advantageous in dealing with low light levels found in forest understorey. We used four independent contrasts of heteroblastic and homoblastic seedlings from the genera Aristotelia, Hoheria, Pseudopanax, and Melicope grown in full-sun (100% sunlight) and shade (5% sunlight) light environments in a glasshouse. The four heteroblastic species had consistently smaller leaves and lower specific leaf area than their paired homoblastic species both in sun and shade. In the shade, there were no consistent differences in leaf anatomy (thickness of leaf blade, cuticle, epidermis, and palisade mesophyll, and stomatal density × stomatal aperture length) or physiology (maximum photosynthetic rate, dark respiration, and light compensation point) between homoblastic and heteroblastic species. However, in the sun, heteroblastic A. fruticosa, P. crassifolius, and M. simplex had appreciably thicker leaf blades as well as higher maximum photosynthetic rates than their homoblastic congeners. These traits suggest heteroblastic seedlings possess leaf traits associated with an advantage in high-light environments. We conclude that the heteroblastic seedling leaf morphology is unlikely to be an adaptation to very low light. Alternative explanations for the functional significance of changing leaf morphology in association with life-stage should be sought.     

Ecological and morphological variation of Canary Island blue tits, Parus caeruleus (Aves: Paridae)

Canary Island blue tits have differentiated within the last million years. Compared with mainland populations they have short wings, large beaks and tarsi, and tend to be less variable in these parameters. Within the archipelago the eastern populations have the shortest wings and tarsi but the widest and deepest beaks. The eastern populations may have been derived from central island ones, following extinction of the original eastern populations during a warm period in the last Ice Age. New measurements have been used to test contrasting predictions of hypotheses that explain inter-island morphological variation in terms of either climatic or ecological influences. Neither hypothesis is sufficient to explain the variation. A model which combines mem is proposed as follows. Mild winter temperatures have selected for small body size and wing length, especially on the hot, dry, eastern islands. On the other pine-forested, islands, selection has favoured relatively long tarsi and long, slender, beaks for efficient foraging among pine foliage. The absence of die pine-dwelling coal tit from the islands has facilitated mis evolutionary shift. It is shown that blue tits in pines forage more in the needles and cones on the Canary Islands than they do on die North African mainland.     

Microarray analysis of vegetative phase change in maize

Vegetative phase change is the developmental transition from the juvenile phase to the adult phase in which a plant becomes competent for sexual reproduction. The gain of ability to flower is often accompanied by changes in patterns of differentiation in newly forming vegetative organs. In maize, juvenile leaves differ from adult leaves in morphology, anatomy and cell wall composition. Whereas the normal sequence of juvenile followed by adult is repeated with every sexual generation, this sequence can be altered in maize by the isolation and culture of the shoot apex from an adult phase plant: an 'adult' meristem so treated reverts to forming juvenile vegetative organs. To begin to unravel the as-yet poorly understood molecular mechanisms underlying phase change in maize, we compared gene expression in two juvenile sample types, leaf 4 and culture-derived leaves 3 or 4, with an adult sample type (leaf 9) using cDNA microarrays. All samples were leaf primordia at plastochron 6. A gene was scored as 'phase induced' if it was up- or downregulated in both juvenile sample types, compared with the adult sample type, with at least a twofold change in gene expression at a P-value of < or =0.005. Some 221 expressed sequence tags (ESTs) were upregulated in juveniles, and 28 ESTs were upregulated in adults. The largest class of juvenile-induced genes was comprised of those involved in photosynthesis, suggesting that maize plants are primed for energy production early in vegetative growth by the developmental induction of photosynthetic genes.     

Temperature sensitivity of nitrogen productivity for Scots pine and Norway spruce

Environmental conditions control physiological processes in plants and thus their growth. The predicted global warming is expected to accelerate tree growth. However, the growth response is a complex function of several processes with both direct and indirect effects. To analyse this problem we have used needle nitrogen productivity, which is an aggregate parameter for production of new foliage. Data on needle dry matter, production, and nitrogen content in needles of Scots pine ( Pinus sylvestris) and Norway spruce ( Picea abies) from a wide range of climatic conditions were collected and needle nitrogen productivities, defined as dry matter production of needles per unit of nitrogen in the needle biomass, were calculated. Our results show that the nitrogen productivity for spruce is insensitive to temperature. However, for pine, temperature affects both the magnitude of nitrogen productivity at low needle biomass and the response to self-shading but the temperature response is small at the high end of needle biomass. For practical applications it may be sufficient to use a species-specific nitrogen productivity that is independent of temperature. Because temperature affects tree growth indirectly as well as through soil processes, the effects of temperature change on tree growth and ecosystem carbon storage should mainly be derived from effects on nitrogen availability through changes in nitrogen mineralization. In addition, this paper summarises data on dry matter, production and nitrogen content of needles of conifers along a temperature gradient.     

叶异形水生植物不同发育阶段叶片的结构和无机碳获取策略

以睡莲(Nymphaea tetragona Georgi)、萍蓬草(Nuphar pumila (Timm) de Candolle)、条叶萍蓬草(Nuphar sagittifolia Pursh)、眼子菜(Potamogeton distinctus A. Bennett)、南方眼子菜(Potamogeton octandrus Poir)和泽泻(Alisma plantago-aquatica L.)6种叶异形水生植物为材料,对它们的叶绿素含量、气孔性状、解剖结构和HCO₃^-利用等指标进行分析,比较了两种不同发育阶段叶片结构和无机碳获取策略的差异。结果显示,幼态叶比成熟叶的叶片更薄,具有更少的细胞层数,幼态叶上下表面均不具备气孔,而成熟叶上表皮有气孔。说明幼态叶在结构上增加细胞比表面积,增加水下吸收无机碳的能力,而成熟叶的结构能更好地吸收大气中的CO₂。pH-drift分析结果证明,幼态叶具有更高获取水下无机碳的能力,以适应沉水环境。本研究还发现眼子菜和南方眼子菜的幼态叶可以利用水体中HCO₃^-为额外碳源,更有利于其在沉水环境中生长。研究结果阐明了叶异形水生植物不同发育阶段叶片结构和利用无机碳获取策略对水下和气生环境的适应性。     

DOES SHADE PROLONG JUVENILE DEVELOPMENT? A MORPHOLOGICAL ANALYSIS OF LEAF SHAPE CHANGES IN CUCURBIT A ARGYROSPERMA SUBSP. SORORIA (CUCURBITACEAE)

Several studies have concluded that shade extends the juvenile phase of plant development based on the prolonged production of juvenile-looking leaves along the shoot. Until now, the alternative hypothesis that leaves produced in shade converge in shape with more juvenile leaves through plastic responses of individual leaves has not been investigated. The literature has shown that differences in shape among leaves in a heteroblastic series are manifest very early in development, often at or near inception, whereas divergence in development between sun and shade leaves does not become apparent until considerably later. This study is the first to distinguish between these alternatives by comparing the developmental morphology of young leaves of the heteroblastic plant Cucurbita argyrosperma subsp. sororia. Differences in shapes of mature leaves along the shoot in sun and shade were quantified in terms of leaf area, perimeter, and shape using truss analysis. Developmental morphology from initiation through expansion was examined for representative transition and for later (adult) leaves using scanning electron microscopy and allometry. Determinants of shape established very early in development were the same for leaves at the same position grown in sun and shade. Differences in morphology between sun and shade leaves at the same position did not arise until these leaves reached lamina lengths greater than 1,000 μm. Thus, the less-lobed, more juvenile looking leaf produced at later positions in the shade arose through later developmental responses of individual leaves to shade, rather than through a prolonged phase of juvenile development.     

Water relations and gas exchange in Coespeletia moritziana (Sch. Bip) Cuatrec., a giant rosette species of the high tropical Andes

Giant rosettes are ones of the most striking features of the vegetation in the high tropical Andes, with Coespeletia moritziana reaching the highest altitudes up to 4,600 m a.s.l. Different from other giant rosettes, this species grows on rock outcrops with poorly developed soils and where water availability may be limited. Two questions are addressed in this study: How does this species respond in terms of water relations to maintain favorable gas-exchange conditions? Considering that adult plants rely on a water-reserving central pith, how do early stages respond to this environment??s extreme conditions? Water relations and gas-exchange studies were carried out on juveniles, intermediate and adult C. moritziana plants during wet and dry seasons in Páramo de Piedras Blancas at 4,200 m a.s.l. Adult plants maintained higher leaf water potentials (??_L) during the wet season, however, no differences between stages were found for the dry season. Minimum dry season ??_L were never near the turgor loss point in any of the stages. Juveniles show a more strict stomatal control during the dry season to maintain a favorable water status. Net photosynthesis significantly decreased in intermediate and juvenile stages from wet to dry seasons. Our results suggest that C. moritziana resists more extreme conditions compared to other Andean giant rosettes     

Diapause in a large aggregation of a tropical beetle

Abstract. 1_˜ Diapause is described for adults of Stenotarsus rotundus Arrow (Endomychidae, Coleoptera) in the tropical lowland forest on Barro Colorado Island, Panama.
2 . Diapause lasted up to 10 months each year, covering some 6 months of the wet season plus 4 months of the dry season. When in diapause the beetles clustered in an aggregation of up to 70 ,000 individuals on the stem of a palm tree, the same individual palm tree in at least four successive years.
3. The habitat and habits of the beetle when not in diapause are unknown, but one or two generations may be produced during this period. Beetles returned to the diapause site from June until late August.
4. Soon after arrival at the diapause site the flight muscles degenerated and did not develop again until February/March. The ovaries were undeveloped when the beetles arrived at the site and remained in that condition until the onset of yolk deposition in March. At the beginning of the rainy season in April the beetles mated and dispersed.
5. Lean dry weight of the beetles did not decrease noticeably during diapause, but fat content did. It is suggested that the beetles do not feed, or feed very little, while in diapause. During diapause, the metabolic rate is surprisingly low considering a mean ambient temperature of 25° C.
6. Juvenile hormone analogue (ZR515) stimulated flight muscle and ovary development. Daylength, when different from LD 12: 12 , initiated a partial flight muscle development, but there was no effect on the ovaries. Field observations suggest that neither temperature nor humidity had an effect on the termination of diapause.
7. Although the development of both flight muscles and ovaries were stimulated by juvenile hormone analogue and both developed around March of each year, it is suggested that the mechanisms causing this development are different.     

Cytokinins and bud morphology in Pinus radiata

Cytokinins (CKs) play essential roles in the regulation of plant growth and development. In the previous paper (Zhang et al. 2001), we reported the detection and identification of a wide spectrum of CKs, including several novel forms, in the buds of Pinus radiata D. Don. In this paper we examine the relationship between the CKs and buds from juvenile and adult trees of P. radiata. During development the morphology of buds alters significantly, from buds bearing primary needles during their juvenile phase to buds sealed in scales at the adult phase. The morphology of adult buds is a very stable character, as fascicle meristems released from apical dominance, or cultured in vitro, produced only secondary needles. However, exogenous CK causes the adult buds to revert to juvenile bud development in vitro . Analyses of the endogenous CKs revealed that juvenile buds had a relatively higher level of isopentenyladenine and isopentenyladenosine, extremely low levels of phosphorylated CKs and a relatively low level of novel CK glycosides. The adult buds contained lower levels of free base and riboside CKs but very high levels of phosphorylated CKs and novel CK glycosides. Possible roles for CKs in the regulation of bud development are discussed.     

EFFECT OF SEED SIZE ON HETEROBLASTIC DEVELOPMENT IN SEEDLINGS OF DESMODIUM PANICULATUM

Seedlings of Desmodium paniculatum derived from different seed sizes differed in heteroblastic development. The transition from juvenile to adult leaves took place at the fourth node in seedlings derived from larger seeds and at the fifth or sixth node in those derived from smaller ones. Cotyledon removal had no effect on this condition, but growth under reduced irradiance delayed the formation of adult leaves in seedlings from larger seeds. Simple and compound leaves had similar unit area photosynthetic rates and stomatal conductance values, but compound leaves had a 30% higher total area. The earlier production of adult leaves would thus give seedlings from larger seeds an advantage in terms of total carbon gain.     

云贵高原抚仙湖近13300年的花粉/炭屑记录*

抚仙湖是云贵高原著名的断陷深水湖,其沉积物蕴藏着流域地质历史时期丰富的环境信息。对钻取自该湖的900cm 湖泊沉积物岩芯进行花粉/炭屑分析及花粉数据的主成分分析表明,抚仙湖流域的植被、气候与火灾在过去的13 300年经历了5个阶段的变化：(1)13 300—10 400cal．a BP,植被以松林为主,伴有山地暗针叶林和常绿阔叶林,表明该时期气候较为冷湿,森林火灾多发,在后期随着温度和湿度的降低,森林火灾愈加频繁。(2)10 400—5 700cal．a BP,松林收缩,常绿阔叶林扩张,出现一定数量的落叶阔叶林,显示该时期气候偏暖偏干;此阶段早期随着气候变暖变干森林火灾的发生延续上阶段高发的状态,直到9 500cal．a BP后随着湿度的增加森林火灾明显减少。(3)5 700—1 800cal．a BP,松林变化较小,常绿/落叶阔叶林比重增大,首次出现了暖热性的枫香林,显示该时期暖湿的气候特征,火灾发生频率低。(4)1 800—500cal．a BP,松林扩张,阔叶林收缩,本阶段后期草本植被比重开始增加,显示该时期气候相对冷干,森林火灾发生频率较高。(5)500cal．a BP至今,松林收缩,落叶阔叶树种增多,草本植物花粉明显增多,显示该时期气候温凉偏干,森林火灾发生频率降低。     

Needle traits of an evergreen, coniferous shrub growing at wind-exposed and protected sites in a mountain region: does Pinus pumila produce needles with greater mass per area under wind-stress conditions?

Snow depth is one of the most important determinants of vegetation, especially in mountainous regions. In such regions, snow depth tends to be low at wind-exposed sites such as ridges, where stand height and productivity are limited by stressful environmental conditions during winter. Siberian dwarf pine ( Pinus pumila Regel) is a dominant species in mountainous regions of Japan. We hypothesized that P. pumila produces needles with greater mass per area at wind-exposed sites than at wind-protected sites because it invests more nitrogen (N) in cell walls at the expense of N investment in the photosynthetic apparatus, resulting in increased photosynthetic N use efficiency (PNUE). Contrary to our hypothesis, plants at wind-exposed site invested less resources in needles, as exhibited by lower biomass, N, Rubisco and cell wall mass per unit area, and had higher photosynthetic capacity, higher PNUE and shorter needle life-span than plants at a wind-protected site. N partitioning was not significantly different between sites. These results suggest that P. pumila at wind-exposed sites produces needles at low cost with high productivity to compensate for a short leaf life-span, which may be imposed by wind stress when needles appear above the snow surface in winter.     

湖南省冷水江市大嵊山大气污染与马尾松衰亡

 作者对湖南省冷水江市大嵊山大气污染与马尾松衰亡进行了研究,研究结果表明大气综合污染指数在3个研究点上分别为1.00(对照)、1.76和2.23。而以林分密度、林分非衰亡率、针叶长度、针叶干重、马尾松平均高、平均胸径等因子计算出的林分外貌综合得分从173.9下降到143.5和132.5。在一年四季,马尾松针叶中叶绿素含量也是清洁区显著高于污染区,而叶绿素a／b值则是污染区的高,峰值较正常的提前。马尾松在受到大气污染作用后,针叶中营养元素含量呈下降趋势,尤其是氮。根据树干解析发现大气污染对马尾松高生长和胸径生长都具有抑制作用,但并不表现出同步效应,对高生长的抑制有一个滞后现象,时间在2～3年。另外,大气污染对马尾松的结果影响很大,污染区的马尾松单株拥有球果数仅为清洁的25％,用灰色系统理论对马尾松材积与相关影响因子进行关联度分析发现,材积与综合污染指数的倒数有很好的关联性,关联度达到了0.773。     

The MADS-box gene DAL1 is a potential mediator of the juvenile-to-adult transition in Norway spruce (Picea abies)

Progression through the plant life cycle involves change in many essential features, most notably in the capacity to reproduce. The transition from a juvenile vegetative and non-reproductive to an adult reproductive phase is gradual and can take many years; in the conifer Norway spruce, Picea abies, typically 20-25 years. We present a detailed analysis of the activities of three regulatory genes with potential roles in this transition in Norway spruce: DAL1, a MADS-box gene related to the AGL6 group of genes from angiosperms, and the two LEAFY-related genes PaLFY and PaNLY. DAL1 activity is initiated in the shoots of juvenile trees at an age of 3-5 years, and then increases with age, whereas both LFY genes are active throughout the juvenile phase. The activity of DAL1 further shows a spatial pattern along the stem of the tree that parallels a similar gradient in physiological and morphological features associated with maturation to the adult phase. Constitutive expression of DAL1 in transgenic Arabidopsis plants caused a dramatic attenuation of both juvenile and adult growth phases; flowers forming immediately after the embryonic phase of development in severely affected plants. Taken together, our results support the notion that DAL1 may have a regulatory role in the juvenile-to-adult transition in Norway spruce.