Generational differences in response to desiccation stress in the desert moss Tortula inermis

BACKGROUND AND AIMS: Active growth in post-embryonic sporophytes of desert mosses is restricted to the cooler, wetter months. However, most desert mosses have perennial gametophytes. It is hypothesized that these life history patterns are due, in part, to a reduced desiccation tolerance for sporophytes relative to gametophytes. METHODS: Gametophytes with attached post-embryonic sporophytes of Tortula inermis (early seta elongation phenophase) were exposed to two levels of desiccation stress, one rapid-dry cycle and two rapid-dry cycles, then moistened and allowed to recover, resume development, and/or regenerate for 35 d in a growth chamber. KEY RESULTS: Gametophytes tolerated the desiccation treatments well, with 93 % survival through regenerated shoot buds and/or protonemata. At the high stress treatment, a significantly higher frequency of burned leaves and browned shoots occurred. Sporophytes were far more sensitive to desiccation stress, with only 23 % surviving after the low desiccation stress treatment, and 3 % surviving after the high desiccation stress treatment. While the timing of protonemal production and sporophytic phenophases was relatively unaffected by desiccation stress, shoots exposed to one rapid-dry cycle produced shoots more rapidly than shoots exposed to two rapid-dry cycles. CONCLUSIONS: It is concluded that sporophytes of Tortula inermis are more sensitive to rapid drying than are maternal gametophytes, and that sporophyte abortion in response to desiccation results from either reduced desiccation tolerance of sporophytes relative to gametophytes, or from a termination of the sporophyte on the part of the gametophyte in response to stress.     

SENSITIVITY OF ANTARCTIC UROSPORA PENICILLIFORMIS (ULOTRICHALES,CHLOROPHYTA) TO ULTRAVIOLET RADIATION IS LIFE‐STAGE DEPENDENT1

The sensitivity of different life stages of the eulittoral green alga Urospora penicilliformis (Roth) Aresch. to ultraviolet radiation (UVR) was examined in the laboratory. Gametophytic filaments and propagules (zoospores and gametes) released from filaments were separately exposed to different fluence of radiation treatments consisting of PAR (P = 400–700 nm), PAR + ultraviolet A (UVA) (PA, UVA = 320–400 nm), and PAR + UVA + ultraviolet B (UVB) (PAB, UVB = 280–320 nm). Photophysiological indices (ETR_max, E_k, and α) derived from rapid light curves were measured in controls, while photosynthetic efficiency and amount of DNA lesions in terms of cyclobutane pyrimidine dimers (CPDs) were measured after exposure to radiation treatments and after recovery in low PAR; pigments of propagules were quantified after exposure treatment only. The photosynthetic conversion efficiency (α) and photosynthetic capacity (rETR_max) were higher in gametophytes compared with the propagules. The propagules were slightly more sensitive to UVB‐induced DNA damage; however, both life stages of the eulittoral inhabiting turf alga were not severely affected by the negative impacts of UVR. Exposure to a maximum of 8 h UVR caused mild effects on the photochemical efficiency of PSII and induced minimal DNA lesions in both the gametophytes and propagules. Pigment concentrations were not significantly different between PAR‐exposed and PAR + UVR–exposed propagules. Our data showed that U. penicilliformis from the Antarctic is rather insensitive to the applied UVR. This amphi‐equatorial species possesses different protective mechanisms that can cope with high UVR in cold‐temperate waters of both hemispheres and in polar regions under conditions of increasing UVR as a consequence of further reduction of stratospheric ozone.     

Chlorophyll concentrations and a/b ratios in mosses collected from exposed and shaded habitats in Kansas

Abstract

Chlorophyll concentrations and a/b ratios of 31 moss species in 16 families collected from various habitats in Kansas were lower than those of most higher vascular plants. Most of the mosses examined had higher chlorophyll concentrations and lower chlorophyll a/b ratios when collected from habitats with low solar irradiance, relative to mosses collected from habitats characterized by high irradiance. This was true for mosses collected from under a forest canopy compared with those collected from exposed sandstone outcrops, and for the same populations of mosses collected before and after forest canopy closure. Such changes in the characteristics of the light-harvesting pigment system appear to represent adaptations enhancing the efficiency of light capture.     

Sporophyte and gametophyte generations differ in their thermotolerance response in the moss Microbryum

BACKGROUND AND AIMS: Actively growing post-embryonic sporophytes of desert mosses are restricted to the cooler, wetter months. However, most desert mosses have perennial gametophytes. It is hypothesized that these life history patterns are due in part to a reduced thermotolerance for sporophytes relative to gametophytes. METHODS: Gametophytes with attached embryonic sporophytes of Microbryum starckeanum were exposed whilst desiccated to thermal episodes of 35 degrees C (1 hr), 55 degrees C (1 hr), 75 degrees C (1 hr) and 75 degrees C (3 hr), then moistened and allowed to recover for 35 d in a growth chamber. KEY RESULTS: All of the gametophytes survived the thermal exposures and produced protonemata, with the majority also producing shoot buds. Symptoms of gametophytic stress (leaf burning and discoloration of entire shoots) were present in lower frequencies in the 55 degrees C exposure. Sporophyte resumption of growth and maturation to meiosis were significantly negatively affected by thermal treatment. Not a single sporophyte exposed to the two higher thermal treatments (75 degrees C for 1 h and 75 degrees C for 3 h) survived to meiosis, and those sporophytes exposed to 75 degrees C that survived to the post-embryonic phenophase took significantly longer to reach this phase. Furthermore, among the thermal treatments where some capsules reached maturity (35 degrees C and 55 degrees C), maternal shoots that produced a meiotic capsule took longer to regenerate through protonemata than maternal shoots aborting their sporophyte, suggestive of a resource trade-off between generations. CONCLUSIONS: Either (1) the inherent sporophyte thermotolerance is quite low even in this desert moss, and/or (2) a gametophytic thermal stress response controls sporophyte viability.     

Towards an understanding of sex determination in Equisetum: an analysis of regeneration in gametophytes of the subgenus Equisetum

Equisetum gametophytes possess extensive powers of regeneration without showing any signs of diminished vigour or abnormal development. Isolated lamellae, pieces of antheridial and archegonial branches were used to analyse gametophyte morphogenesis and sex determination. Intact individuals and subcultured sexual branches never revert to a vegetative state. Transplantation of apical pieces of archegonial branches on to fresh medium delays the onset of antheridium formation indicating phenotypic control. On the other hand, male branches invariably continue producing antheridia. Experimentally induced adventitious branches on lamellae are interpreted as the result of physiological isolation from inhibitors. In intact individuals these adventitious branches are always antheridial: after subculturing, archegonial branches are also produced, demonstrating that the gametophytes of Equisetum are all potentially bisexual. Isolated lamellae regenerate secondary gametophytes, the sexual ratios between which vary between species, but lamellae from archegonial tissues invariably produce more female individuals, probably reflecting the different sizes of the respective lamellae. The view that mixtures of intact male and bisexual Equisetum gametophytes result from inherent physiological differences between the spores is not supported.     

Sporophytes and gametophytes of Dicranaceae from the Santonian (Late Cretaceous) of Georgia, USA

A new species (Campylopodium allonense) of the moss family Dicranaceae is described for fossil sporophyte capsules and associated gametophytes from the late Santonian (Late Cretaceous) Buffalo Creek Member of the Gaillard Formation in central Georgia, USA. The sporophyte capsules are most comparable to those of the living genus Campylopodium. Sporangia are curved, cylindrical, and strumose, with an obliquely rostrate operculum, cucullate calyptra, and compound annulus. The peristome is haplolepidous with 16 dicranoid, apically bifid teeth that are vertically striate on the outer surface and asymmetrically trabeculate on the inner surface. Spores are spherical, alete, and finely rugose, and thus differ from the finely papillose spores of extant Campylopodium. Associated fossil gametophytes are consistent with the morphology of extant Campylopodium and have leaves with a broad sheathing base and a narrow blade. Spores identical to those in the sporangium occur on the leaf surfaces of one of the gametophyte specimens, providing circumstantial evidence that both sporophyte and gametophyte belong to the same species. Inadequacies of the moss fossil record have led to contrasting interpretations of the timing of evolutionary change in this lineage since the Paleozoic. Campylopodium allonense unequivocally provides the earliest evidence of Dicranaceae in the fossil record. This material, along with other fossil mosses from this late Santonian locality, indicates the presence of modern families of mosses in the Cretaceous. In a phylogenetic context, these fossils from two different subclasses imply that mosses were already diverse by the Late Cretaceous.     

Differential responses of tetrasporophytes and gametophytes of Mazzaella laminarioides (Gigartinales,Rhodophyta) under solar UV radiation

The effects of solar UV radiation on mycosporine‐like amino acids (MAAs), growth, photosynthetic pigments (Chl a, phycobiliproteins), soluble proteins (SP), and C and N content of Mazzaella laminarioides tetrasporophytes and gametophytes were investigated. Apical segments of tetrasporophytes and gametophytes were exposed to solar radiation under three treatments (PAR [P], PAR+UVA [PA], and PAR+UVA+UVB [PAB]) during 18 d in spring 2009, Punta Arenas, Chile. Samples were taken after 2, 6, 12, and 18 d of solar radiation exposure. Most of the parameters assessed on M. laminarioides were significantly influenced by the radiation treatment, and both gametophytes and tetrasporophytes seemed to respond differently when exposed to high UV radiation. The two main effects promoted by UV radiation were: (i) higher synthesis of MAAs in gametophytes than tetrasporophytes at 2 d, and (ii) a decrease in phycoerythrin, phycocyanin, and SPs, but an increase in MAA content in tetrasporophytes at 6 and 12 d of culture. Despite some changes that were observed in biochemical parameters in both tetrasporophytes and gametophytes of M. laminarioides when exposed to UVB radiation, these changes did not promote deleterious effects that might interfere with the growth in the long term (18 d). The tolerance and resistance of M. laminarioides to higher UV irradiance were expected, as this intertidal species is exposed to variation in solar radiation, especially during low tide.     

Arbuscular mycorrhizal structure and fungi associated with mosses

We investigated the colonization and diversity of arbuscular mycorrhizal (AM) fungi associated with 24 moss species belonging to 16 families in China. AM fungal structures, i.e. spores, vesicles, hyphal coils (including intracellular hyphae), or intercellular nonseptate hyphae, were found in 21 moss species. AM fungal structures (vesicles, hyphal coils, and intercellular nonseptate hyphae) were present in tissues of 14 moss species, and spores and nonseptate hyphae on the surface of gametophytes occurred in 15 species. AM fungal structures were present in 11 of the 12 saxicolous moss species and in six of the ten terricolous moss species, but absent in two epixylous moss species. AM fungal structures were only observed in moss stem and leaf tissues, but not in rhizoids. A total of 15 AM fungal taxa were isolated based on trap culture with clover, using 13 moss species as inocula. Of these AM fungi, 11 belonged to Glomus, two to Acaulospora, one to Gigaspora, and one to Paraglomus. Our results suggest that AM fungal structures commonly occur in most mosses and that diverse AM fungi, particularly Glomus species, are associated with mosses.     

Effects of Different Photoperiods and Chilling Treatments on Growth and Reproduction of Cardamine flexuosa With. and Its Ecological Implications

Abstract Phenotypic variability of Cardamine flexuosa (Cruciferae) was examined in response to different lengths of exposure to low temperature (5°C) at the juvenile stage, and to two photoperiod regimes (8 and 16 hrs day-lengths) in the subsequent growth period. The results indicated that this species had a facultative chilling or long-day requirement for flowering. The long-day and chilling treatments both caused an earlier onset of stem internode elongation. Longer chilling treatments reduced the number of nodes and increased internode length under both long and short photoperiod regimes. Prolonged chilling treatments followed by long-day photoperiod remarkably induced more numerous basal branches and inflorescences, as a result producing more siliques. Remarkable rosette leaves Were formed at the base of the main stem when partial chilling and the short-day treatment suppressed stem internode elongation. The responses to chilling and long-day treatment closely resembled the synchronized flowering of this species in spring in the field. Delayed flowering under the short-day treatment resembled size- or age-dependent flowering in late summer to autumn in the field populations.     

Effects of elevation, wave exposure, and year on the proportion of gametophytes and tetrasporophytes in Mazzaella parksii (Rhodophyta, Gigartinaceae) populations

We investigated the effects of elevation, wave exposure, and year on the proportion of gametophytes and tetrasporophytes in populations of the intertidal red seaweed Mazzaella parksii (= M. cornucopiae, Gigartinaceae) from the Pacific coast of Canada. In July 2002, we determined the life-history phase of 864 frond samples from four populations from Barkley Sound, using the resorcinol-acetal test. For these populations, gametophytes were significantly more numerous than tetrasporophytes, and the proportion of gametophytes was significantly higher at higher elevations. The proportion of gametophytes varied directly (although less markedly) with the degree of wave exposure, but significance could not be assessed for this factor. All of these patterns were spatially consistent, as they held for two different rocky points with similar physical characteristics. One of these points had also been surveyed in 1994-1995, which allows us to conclude that the population-wise gametophyte predominance and the positive relationship between elevation and the proportion of gametophytes are stable features of this species in this area. However, the average proportion of gametophytes was similar between the studied levels of wave exposure in 1994–1995; such a difference revealed as non-significant when data were combined with those for 2002. This suggests that wave exposure is not important in determining the proportion of life-history phases for this species. Comparisons with other species are done in search of general patterns for the Gigartinaceae.     

GROWTH REGULATION BY ETHYLENE IN FERN GAMETOPHYTES. I. EFFECTS ON PROTONEMAL AND RHIZOIDAL GROWTH AND INTERACTION WITH AUXIN

Three responses resulted from the treatment with ethylene of dark-grown gametophytes of Onoclea sensibilis. Elongation of the filament was increased, elongation of the rhizoid was decreased, and cell division was inhibited. The optimal ethylene concentrations were between 0.01 and 0.1 ppm. Filament elongation was very tolerant of high ethylene concentrations, since up to 1,000 ppm did not inhibit growth below control levels. Enclosing cultures of gametophytes in chambers of limited volumes produced all the effects of treatment with ethylene, but the responses to sealing were eliminated if plants were enclosed in a chamber which contained a solution of mercuric Perchlorate. This evidence that gametophytes produced ethylene was substantiated by a direct gas chromatographic demonstration of ethylene formation. The growth-regulating effects of ethylene and auxin appeared to be independent in filament and rhizoid growth. Inhibition of elongation by supra-optimal auxin concentrations could not be attributed to an auxin-stimulated production of ethylene.     

EFFECTS OF ACIDIC SOLUTIONS ON SEXUAL REPRODUCTION OF PTERIDIUM AQUILINUM

Experiments were performed to determine the effects of acidic solutions on spermatozoid motility and fertilization of gametophytes of Pteridium aquilinum. Buffered solutions (0.0025 m ) were used to simulate exposures to acidic precipitation for up to a 3.5 hr exposure. Experimental results suggest that the spermatozoid population can be subdivided into several groups with respect to pH sensitivity; about 25% spermatozoids are immobile one min after exposure to pH 6.1 buffer while about an equal percentage remain motile after 30 min exposure to buffer of pH 5.1. Between these two response extremes are two other subpopulations. One is quite sensitive to pH but shows some recovery if pH is between 5.6 and 6.1, while the second subpopulation does not seem to exhibit any motility recovery at all but is more resistant to acidity than the first subpopulation. To complement experiments that evaluate spermatozoid responses, experiments were performed to view the process of fertilization under controlled environmental conditions as well as under the canopy of a forest. Fertilization of gametophytes in uncovered petri dishes under a forest canopy was similar to results in aseptic culture after gametophytes were exposed to various pH levels and 86.6 μM sulfate. Although there were some differences between results obtained under aseptic culture conditions and cultures maintained under a forest canopy, it is evident that a lower buffer pH decreased fertilization. Fertilization at pH 4.5 and 3.6 was about one-half that occurring at pH 6.1. Fertilization in gametophytes exposed to pH 3.0 was about 10-20% of that occurring at pH 6.1. Addition of 86.6 μM sulfate decreased fertilization under all culture conditions. These experimental results suggest that fertilization in P. aquilinum may be used as a bioindicator of contaminants in rainwater. The results demonstrate that spermatozoid motility (and the process of fertilization) is more acid sensitive than gametophytic and sporophytic tissues.     

Differences in growth trajectory and strategy of two sympatric congeneric species in an Indonesian floodplain forest

Whole-plant development trajectories and sapling leaf displays were compared for two sympatric congeneric species, Pterospermum diversifolium and P. javanicum, in a tropical floodplain forest in East Kalimantan, Indonesia. We assessed their growth strategies and developed hypotheses for their coexistence within the community. Pterospermum diversifolium retains a monoaxial growth habit that promotes quick stem elongation; thus, it is taller when branches are initiated than is P. javanicum. The species differed significantly in height growth and total crown expansion per unit increment of biomass: monoaxial P. diversifolium saplings devote more effort to stem elongation, whereas branched P. javanicum saplings devote more effort to branch expansion. Monoaxial P. diversifolium sustained more severe self-shading than P. javanicum. The sapling growth strategy of P. diversifolium appears to be dynamic, emphasizing the opportunistic use of light following a disturbance, whereas that of P. javanicum appears to be static, optimizing leaf display for current light conditions. The advantages of these strategies depend on context, and the two species may coexist within a community by adopting different regeneration niches based on differing understory light conditions: P. diversifolium is favored over P. javanicum at high light levels, but the opposite is true at low light levels.     

Testing adaptive plasticity to UV: costs and benefits of stem elongation and light-induced phenolics

On exposure to ultraviolet radiation (UV), many plant species both reduce stem elongation and increase production of phenolic compounds that absorb in the UV region of the spectrum. To demonstrate that such developmental plasticity to UV is adaptive, it is necessary to show that the induced phenotype is both beneficial in inductive environments and maladaptive in non-inductive environments. We measured selection on stem elongation and phenolic content of seedlings of Impatiens capensis transplanted into ambient-UV and UV-removal treatments. We extended the range of phenotypes expressed, and thus the opportunity for selection in each UV treatment, by pretreating seedlings with either a low ratio of red:far-red wavelengths (R:FR), which induced stem elongation and reduced phenolic concentrations, or high R:FR, which had the opposite effect on these two phenotypic traits. Reduced stem length relative to biomass was advantageous for elongated plants under ambient UV, whereas increased elongation was favored in the UV-removal treatment. Selection favored an increase in the level of phenolics induced by UV in the ambient-UV treatment, but a decrease in phenolics in the absence of UV. These results are consistent with the hypotheses that reduced elongation and increased phenolic concentrations serve a UV-protective function and provide the first explicit demonstration in a wild species that plasticity of these traits to UV is adaptive. The observed cost to phenolics in the absence of UV may explain why many species plastically upregulate phenolic production when exposed to UV, rather than evolve constitutively high levels of these compounds. Finally, pretreatment with low R:FR simulating foliar shade did not exacerbate the fitness impact of UV exposure when plants had several weeks to acclimate to UV. This observation suggests that the evolution of adaptive shade avoidance responses to low R:FR in crowded stands will not be constrained by increased sensitivity to UV in elongated plants when they overtop their neighbors.     

Bud and crown architecture of white spruce and black spruce

Needle primordia in buds and branch lengths were assessed in the crown of a plantation-grown white spruce tree. There was a gradation in needle primordia in buds in branches within the crown. The largest number of primordia was in the terminal bud of the leading main stem shoot, with the number in first-order whorl lateral shoot terminal buds decreasing from whorl 1 to whorl 4, below which buds contained a similar small number of primordia (about one-third as many as in the terminal shoot). Previous year's shoot elongation followed a similar pattern (i.e., elongation of whorl branches was greater closer to the top of the tree and elongation in the fourth through ninth whorls was about one-third that of the main stem leader). Higher order branches within whorls had within-branch gradation in shoot elongation and number of needle primordia, with older branches having as few as 16–30 primordia in buds and 3–4 cm elongation for high-order branches on older main stem whorls. There were strong correlations between the number of primordia in branch terminal buds and branch length/diameter and bud length/diameter/volume. In both black spruce and white spruce, there were strong correlations of number of needle primordia in main stem leader terminal buds with number of needle primordia in terminal buds of first and second whorl leaders.     

Development of the leafy shoot in Sphagnum (Bryophyta) involves the activity of both apical and subapical meristems

This light- and electron-microscope study of four species of Sphagnum reveals that stem elongation involves meristematic activities unique to the group and hitherto unrecognized. The internal tissue of the mature stem arises by the concerted activity of an apical (primary) and a subapical (secondary) meristem. The primary meristem comprises the immediate derivatives of the single apical cell. Following a small number of divisions, the primary derivatives differentiate into highly vacuolate parenchymatous cells with a storied arrangement. Subsequently, the large vacuoles are replaced by numerous small vacuoles and the cells then divide repeatedly, by transverse septa, producing files of about nine short cells. Finally, ninefold elongation of these secondary cells is responsible for extension growth of the main stem below the mature capitulum. An early step in primary differentiation is the confinement of pre-existing plasmodesmata to distinct pitted areas. Further enlargement of the cells during primary and secondary differentiation involves the thickening of non-pitted wall areas, followed by expansion and thinning out, while the pitted areas remain virtually unchanged. A cortical array of microtubules is regularly found in association with non-pitted wall areas, while the unexpanded pitted areas are associated with smooth endoplasmic reticulum showing continuity with desmotubules. Though sharing much the same cytology as the conducting cells in bryoid mosses, in terms of their development the central stem cells in Sphagnum are not homologous with those of other mosses. The unique mode of stem development may be an important factor in the ecological success of Sphagnum .     

Metallothionein responses in the Asiatic clam (Corbicula fluminea) after exposure to trivalent arsenic

The main objective of this work was to evaluate arsenic effects on metallothionein (MT) induction by exposing a freshwater Asiatic clam (Corbicula fluminea) to different concentrations of this metalloid. The presence of MT-like proteins was detected by sodium dodecyl sulfate polyacrylamide gel electrophoresis and compared with a standard rabbit MT. In addition, the polarographic response showed good correspondence between standard MT and MT-like curves from C. fluminea, allowing MT quantification. The results show that clams exposed to different concentrations of arsenic are able to induce significant levels of MTs. Although variability was found in MT induction, significant differences in MT levels were found after 28 days of exposure in all treatments in comparison with the controls, suggesting that exposure to arsenic induced MT-like proteins in C. fluminea.     

The Use of Mixtures of Sulfur and Bracken Litter to Reduce pH of Former Arable Soils and Control Ruderal Species

High soil pH has been highlighted as a constraint to the restoration of heathland on ex‐arable land. Previous studies at the Minsmere Reserve of the Royal Society for the Protection of Birds (RSPB) in England have shown that it is possible to acidify ex‐arable soils using elemental sulfur and bracken litter, although sulfur (S) is more effective. Current recommendations suggest that 4 tS/ha need to be applied to reduce soil pH below pH 4, control vigorous ruderal species, and create conditions suitable for Calluna vulgaris (heather) establishment. However, S is relatively expensive, and as bracken litter is moderately abundant within the reserve, it made economic sense to evaluate the potential for mixing S with bracken to see if adequate pH reductions could be achieved at lower S rates. Accordingly an experiment was designed to test the effects of combining S (0–8 t S/ha) and bracken litter (0–10 cm depth layers) on (1) soil pH, (2) cover of ruderal species, and (3) the developing plant community. Significant interactions were detected, especially in the period immediately after application. Where bracken litter was applied the soil pH fell immediately; in contrast, S took at least six months to start reducing pH. Where mixtures were applied there was a synergistic effect, which produced a lower pH than the S or bracken litter applied alone. These effects were most marked at low S application rates, between 0.5–4 t S/ha. The effects of the bracken litter addition also reduced the growth of ruderal species in the period immediately after application, probably through a combination of acidification and physical smothering. There is, therefore, a clear potential to acidify ex‐arable soils using combinations of S and bracken litter in schemes designed to restore Calluna heathland.     

Influence of annually fluctuating seed-cone production and climatic factors on the upper-crown expansion of canopy trees of Abies mariesii Masters (Pinaceae)

To understand the influence of seed-cone production on the upper-crown expansion of canopy trees of Abies mariesii Masters, the influence of the annually fluctuating number of seed cones on the terminal-leader length of the main stem and primary branches was analyzed using seed-cone rachises remaining on the terminal leaders of the primary branches. The influence of climatic factors during the period of vegetative-bud growth and shoot elongation was also analyzed. For the upper-crown expansion, factors influencing the number of horizontally elongating branches on the top of the main stem were also examined. According to model fitting, the terminal-leader length of the main stem was negatively influenced by seed-cone production 1 year before shoot elongation. In contrast, the terminal-leader length of the primary branches was not clearly influenced by seed-cone production, but was positively influenced by the mean temperature in September 1 year before shoot elongation and in July in the year of shoot elongation. The number of horizontally elongating branches on the top was positively influenced by the terminal-leader length of the main stem 1 year before horizontal elongation on the top and negatively influenced by seed-cone production 2 years earlier. Thus, in the upper part of the crown, the allocation of photosynthate to seed cones mainly affects the arrangement of needles and seed cones up to the higher positions. This trade-off between seed-cone production and upper-crown expansion influences the canopy shape and reproductive strategy of A. mariesii .