A molecular module connects abscisic acid with auxin signals to facilitate seasonal wood formation in Populus.

Perennial trees have a recurring annual cycle of wood formation in response to environmental fluctuations. However, the precise molecular mechanisms that regulate the seasonal formation of wood remain poorly understood. Our prior study indicates that VCM1 and VCM2 play a vital role in regulating the activity of the vascular cambium by controlling the auxin homoeostasis of the cambium zone in Populus. This study indicates that abscisic acid (ABA) affects the expression of VCM1 and VCM2, which display seasonal fluctuations in relation to photoperiod changes. ABA-responsive transcription factors AREB4 and AREB13, which are predominantly expressed in stem secondary vascular tissue, bind to VCM1 and VCM2 promoters to induce their expression. Seasonal changes in the photoperiod affect the ABA amount, which is linked to auxin-regulated cambium activity via the functions of VCM1 and VCM2. Thus, the study reveals that AREB4/AREB13-VCM1/VCM2-PIN5b acts as a molecular module connecting ABA and auxin signals to control vascular cambium activity in seasonal wood formation.

Phosphorus addition regulates the growth of Chinese fir by changing needle nitrogen fractions in growing and dormant seasons.

Forest productivity is generally limited by nutrient scarcity. This study aims to reveal seasonal interactions among leaf carbon (C), nitrogen (N) fractions and tree growth driven by nutrient addition in a subtropical forest. Here, a field nutrient addition experiment was conducted with six treatments, namely, +N5 (5 g N m-2 yr-1), +N10 (10 g N m-2 yr-1), +P5 (5 g P m-2 yr-1), +N5 + P5, +N10 + P5, and control (N0 + P0). C fractions (structural and non-structural carbohydrates) and N fractions (soluble N, nucleic N and protein N) in needles as well as tree growth indicated by basal area increment (BAI) were measured in growing and dormant seasons. Total N and protein N in old needles were significantly increased by P addition, while no significant differences of non-structural carbohydrates in young (<1-year old) and old needles (>1-year old) were detected among the treatments in both seasons. N and P addition increased the structural carbohydrates of old needles in dormant season. P addition decreased and increased tree growth in growing and dormant seasons, respectively. The variation of BAI was explained 18.3 % by total N and 17.8 % by protein N in growing season, and was explained 33.9 % by total N and 34.2 % by protein N in dormant season. Our study suggested that the P addition effect on Chinese fir growth mostly depends on needle N fractions. This study highlights tree seasonal growth driven by nutrient alteration might be characterized by leaf N fractions rather than C fractions in subtropical forests.

When to branch: seasonal control of shoot architecture in trees.

Long-lived perennial plants optimize their shoot architecture by responding to seasonal cues. The main strategy used by plants of temperate and boreal regions with respect to surviving the extremely unfavourable conditions of winter comprises the protection of their apical and lateral meristematic tissues. This involves myriads of transcriptional, translational and metabolic changes in the plants because shoot architecture is controlled by multiple pathways that regulate processes such as bud formation and flowering, small RNAs, environmental factors (especially light quality, photoperiod and temperature), hormones, and sugars. Recent studies have begun to reveal how these pathways are recruited for the seasonal adaptation and regulation of shoot architecture in perennial plants, including the role of a regulatory module consisting of antagonistic players terminal flower 1 (TFL1) and like-ap1 (LAP1) in the hybrid aspen. Here, we review recent progress in our understanding of the genetic control of shoot architecture in perennials compared to in annuals.

Seasonal Growth of the Purple Sea Urchin Heliocidaris crassispina Revealed by Sequential Fluorochrome Tagging.

Many studies have applied fluorochrome tagging to examine the growth of animals with calcified skeletons, but most of them have used only a single tag to determine the annual growth rate. We used sequential fluorochrome tagging to study the seasonal growth of the purple sea urchin Heliocidaris crassispina in Hong Kong waters from February 2012 to February 2013. Sea urchins ranging from 18.9 to 42.7 mm in test diameter had a yearly growth from 0.6 to 13.0 mm. During that year, the sea urchins grew from 0.6 to 5.0 mm in test diameter during the first six months, and from 0.4 to 10.2 mm in test diameter in the second six months. The seasonal differences in growth were confirmed using the von Bertalanffy model. The growth was clear for young sea urchins, especially for individuals less than 5 years old, but was not evident for sea urchins older than 7 years. The seasonal differences in growth were probably related to the reproductive cycle and the seasonal differences in environmental conditions. Our empirical results provide the first evidence of seasonal changes in growth for H. crassispina, demonstrating the usefulness of sequential fluorochrome tagging in studying the growth of sea urchins in the field. We also identify the problem of low recovery of tagged individuals and provide recommendations to improve the tagging procedure.

Large variations in eutrophication among estuaries reflect massive differences in composition and biomass of macroalgal drift.

The characteristics of detached macroalgae (drift) in nearby highly eutrophic and mesotrophic estuaries in south-western Australia are compared to elucidate the magnitude and types of changes that occur in macroalgal drift when estuaries receive excessive nutrient input. Drift characteristics in the large basins of the microtidal, eutrophic Peel-Harvey and mesotrophic Swan-Canning, which is not subjected to large nutrient inputs directly from agricultural land, differed markedly. Biomass (dry weight) in mesotrophic estuary was dominated by rhodophytes (92%), particularly Laurencia and Hypnea, and in eutrophic estuary by opportunistic chlorophytes (68%), especially Chaetomorpha and Ulva. Prevalence and biomass of drift were far greater in the eutrophic estuary, particularly during summer and autumn when macroalgal growth rose sharply. Macroalgal biomass in the eutrophic estuary was positively related to salinity. These results facilitate predictions of how climatic and other anthropogenic changes influence extent of macroalgal growth and thus change the estuarine environment.

Timing of spring xylogenesis in temperate deciduous tree species relates to tree growth characteristics and previous autumn phenology.

We explored the timing of spring xylogenesis and its potential drivers in homogeneous mature forest stands in a temperate European region. Three species with contrasting leaf development dynamics and wood anatomy were studied: European beech, silver birch and pedunculate oak. Detailed phenological observations of xylogenesis and leaf phenology were performed from summer 2017 until spring 2018. Cambium reactivation (CR) occurred before the buds of oak and birch were swollen, whereas these two phenological phases were concurrent for beech. On the other hand, initial earlywood vessels were fully differentiated (FDIEV) after leaf unfolding for all three species. Timing of CR was correlated to average ring-width of the last 10 years (2008-17), tree diameter and, partially, with tree age. In addition, the timing of FDIEV was correlated to tree age and previous year's autumn phenology, i.e., timing of wood growth cessation and onset of leaf senescence. Multivariate models could explain up to 68% of the variability of CR and 55% of the variability of FDIEV. In addition to the 'species' factor, the variability could be explained by ca 30% by tree characteristics and previous year's autumn phenology for both CR and FDIEV. These findings are important to better identify which factors (other than environment) can be driving the onset of the growing season, and highlight the influence of tree growth characteristics and previous year's phenology on spring wood phenology, wood formation and, potentially, forest production.

A genetic framework for regulation and seasonal adaptation of shoot architecture in hybrid aspen.

Shoot architecture is critical for optimizing plant adaptation and productivity. In contrast with annuals, branching in perennials native to temperate and boreal regions must be coordinated with seasonal growth cycles. How branching is coordinated with seasonal growth is poorly understood. We identified key components of the genetic network that controls branching and its regulation by seasonal cues in the model tree hybrid aspen. Our results demonstrate that branching and its control by seasonal cues is mediated by mutually antagonistic action of aspen orthologs of the flowering regulators TERMINAL FLOWER 1 (TFL1) and APETALA1 (LIKE APETALA 1/LAP1). LAP1 promotes branching through local action in axillary buds. LAP1 acts in a cytokinin-dependent manner, stimulating expression of the cell-cycle regulator AIL1 and suppressing BRANCHED1 expression to promote branching. Short photoperiod and low temperature, the major seasonal cues heralding winter, suppress branching by simultaneous activation of TFL1 and repression of the LAP1 pathway. Our results thus reveal the genetic network mediating control of branching and its regulation by environmental cues facilitating integration of branching with seasonal growth control in perennial trees.

Branching Regulator BRC1 Mediates Photoperiodic Control of Seasonal Growth in Hybrid Aspen.

Cessation of growth as winter approaches is a key adaptive trait for survival of perennial plants, such as long-lived trees native to boreal and temperate regions [1, 2]. The timing of growth cessation in these plants is controlled by photoperiodic cues. As shown recently, perception of growth-repressive short photoperiod (SP) mediated via components of circadian clock results in downregulation of the tree ortholog of Arabidopsis flowering regulator FLOWERING LOCUS T (FT), FT2 [3, 4]. Downregulation of FT2 results in suppression of downstream components LAP1 (orthologous to the Arabidopsis floral meristem identity gene APETALA1) and AIL1 (orthologous to AINTEGUMENTA in Arabidopsis), culminating in induction of growth cessation and bud set [5-7]. Results presented here reveal that, in addition to the CO/FT pathway, a photoperiodically controlled negative feedback loop involving a tree ortholog of Arabidopsis BRANCHED1 (BRC1) (a member of TEOSINTE BRANCHED 1, CYCLOIDEA, PCF family), LAP1, and FT2 participates in regulation of seasonal growth in the model tree hybrid aspen. In growth-promotive long photoperiod, LAP1 suppresses expression of BRC1, but upon perception of growth-repressive SP, downregulation of LAP1 de-represses expression of its downstream target BRC1. BRC1 physically interacts with FT2, and BRC1-FT interaction further reinforces the effect of SP and triggers growth cessation by antagonizing FT action. Accordingly, BRC1 gain and loss of function result in early and retarded growth cessation responses to SP, respectively. Thus, these results reveal a regulatory feedback loop that reinforces responses to SP and induction of seasonal growth cessation.

Comparison of Growth Rate and Nutrient Content of Five Microalgae Species Cultivated in Greenhouses.

The effect of different environmental conditions on the growth rate, biomass production, nutrient composition, and phenolic content of the microalgae species Chlorella vulgaris, Botryococcus braunii, Chlamydomonas reinhardtii, Euglena gracilis, and Nannochloropsis oculata was investigated. The experiments were conducted in open bioreactors in a greenhouse in three different periods (during October, March, and June), and in a controlled environment in a closed plant growth chamber. It was found that the growth rate and production of C. vulgaris and B. braunii was higher during March, C. reinhardtii and N. oculata grew better in June, and the growth of E. gracilis was similar in March and June. The lipid content of the biomass of all five species increased with increasing light intensity and temperature, while the nitrogen free extractable (NFE) content decreased and the protein, fiber, moisture, and ash content remained relatively unaffected. The phenolic content varied from species to species with E. gracilis having the highest and N. oculata the lowest content among the species studied. The results can be taken into account when cultivating the different microalgae studied in full scale applications, such as in open raceway bioreactors, where conditions could be adjusted to obtain the most favorable growth conditions, depending on the particular species cultivated.

Lianas have a seasonal growth advantage over co-occurring trees.

The seasonal growth advantage hypothesis posits that plant species that grow well during seasonal drought will increase in abundance in forests with increasing seasonality of rainfall both in absolute numbers and also relative to co-occurring plant species that grow poorly during seasonal drought. That is, seasonal drought will give some plant species a growth advantage that they lack in aseasonal forests, thus allowing them attain higher abundance. For tropical forest plants, the seasonal growth advantage hypothesis may explain the distribution of drought-adapted species across large-scale gradients of rainfall and seasonality. We tested the seasonal growth advantage hypothesis with lianas and trees in a seasonal tropical forest in central Panama. We measured the dry-season and wet-season diameter growth of 1,117 canopy trees and 648 canopy lianas from 2011 to 2016. We also evaluated how lianas and trees responded to the 2015-2016 El Niño, which was the third strongest el Niño drought on record in Panama. We found that liana growth rate was considerably higher during the dry-season months than the wet-season months in each of the five years. Lianas achieved one-half of their annual growth during the 4-month dry season. By contrast, trees grew far more during the wet season; they realized only one-quarter of their annual growth during the dry season. During the strong 2015-2016 El Niño dry season, trees essentially stopped growing, whereas lianas grew unimpeded and as well as during any of the previous four dry seasons. Our findings support the hypothesis that seasonal growth gives lianas a decided growth advantage over trees in seasonal forests compared to aseasonal forests, and may explain why lianas peak in both absolute and relative abundance in highly seasonal tropical forests. Furthermore, the ability of lianas to grow during a strong el Niño drought suggests that lianas will benefit from the predicted increasing drought severity, whereas trees will suffer, and thus lianas are predicted to increase in relative abundance in seasonal tropical forests.

Photoperiodic Regulation of Shoot Apical Growth in Poplar.

Woody perennials adapt their genetic traits to local climate conditions. Day length plays an essential role in the seasonal growth of poplar trees. When photoperiod falls below a given critical day length, poplars undergo growth cessation and bud set. A leaf-localized mechanism of photoperiod measurement triggers the transcriptional modulation of a long distance signaling molecule, FLOWERING LOCUS T (FT). This molecule targets meristem function giving rise to these seasonal responses. Studies over the past decade have identified conserved orthologous genes involved in photoperiodic flowering in Arabidopsis that regulate poplar vegetative growth. However, phenological and molecular examination of key photoperiod signaling molecules reveals functional differences between these two plant model systems suggesting alternative components and/or regulatory mechanisms operating during poplar vegetative growth. Here, we review current knowledge and provide new data regarding the molecular components of the photoperiod measuring mechanism that regulates annual growth in poplar focusing on main achievements and new perspectives.

A modified von Bertalanffy growth model dependent on temperature and body size.

Fish growth models are widely used in fisheries as well in aquacultures and ecology. Water temperature is one of the most important factors determining the growth of fish. In the present study, we propose a growth model that includes the effect of water temperature on the growth in the von Bertalanffy growth model. Our model was applied to fit the growth data of bullhead (Cottus gobio), brown trout (Salmo trutta L.), juvenile salmon (Salmo salar), and Araucanian herring (Strangomera bentincki). The model reproduces the growth patterns of each species and fits a set of appropriate parameter values for each species. Moreover, the model reflects the seasonal growth rates quite well.

Making the most of survey data: Incorporating age uncertainty when fitting growth parameters.

Individual growth is an important parameter and is linked to a number of other biological processes. It is commonly modeled using the von Bertalanffy growth function (VBGF), which is regularly fitted to age data where the ages of the animals are not known exactly but are binned into yearly age groups, such as fish survey data. Current methods of fitting the VBGF to these data treat all the binned ages as the actual ages. We present a new VBGF model that combines data from multiple surveys and allows the actual age of an animal to be inferred. By fitting to survey data for Atlantic herring (Clupea harengus) and Atlantic cod (Gadus morhua), we compare our model with two other ways of combining data from multiple surveys but where the ages are as reported in the survey data. We use the fitted parameters as inputs into a yield-per-recruit model to see what would happen to advice given to management. We found that each of the ways of combining the data leads to different parameter estimates for the VBGF and advice for policymakers. Our model fitted to the data better than either of the other models and also reduced the uncertainty in the parameter estimates and models used to inform management. Our model is a robust way of fitting the VBGF and can be used to combine data from multiple sources. The model is general enough to fit other growth curves for any taxon when the age of individuals is binned into groups.

Photoperiod- and temperature-mediated control of growth cessation and dormancy in trees: a molecular perspective.

Background: How plants adapt their developmental patterns to regular seasonal changes is an important question in biology. The annual growth cycle in perennial long-lived trees is yet another example of how plants can adapt to seasonal changes. The two main signals that plants rely on to respond to seasonal changes are photoperiod and temperature, and these signals have critical roles in the temporal regulation of the annual growth cycle of trees. Scope: This review presents the latest findings to provide insight into the molecular mechanisms that underlie how photoperiodic and temperature signals regulate seasonal growth in trees. Conclusion: The results point to a high level of conservation in the signalling pathways that mediate photoperiodic control of seasonal growth in trees and flowering in annual plants such as arabidopsis. Furthermore, the data indicate that symplastic communication may mediate certain aspects of seasonal growth. Although considerable insight into the control of phenology in model plants such as poplar and spruce has been obtained, the future challenge is extending these studies to other, non-model trees.

Dual role of tree florigen activation complex component FD in photoperiodic growth control and adaptive response pathways.

A complex consisting of evolutionarily conserved FD, flowering locus T (FT) proteins is a regulator of floral transition. Intriguingly, FT orthologs are also implicated in developmental transitions distinct from flowering, such as photoperiodic control of bulbing in onions, potato tuberization, and growth cessation in trees. However, whether an FT-FD complex participates in these transitions and, if so, its mode of action, are unknown. We identified two closely related FD homologs, FD-like 1 (FDL1) and FD-like 2 (FDL2), in the model tree hybrid aspen. Using gain of function and RNAi-suppressed FDL1 and FDL2 transgenic plants, we show that FDL1 and FDL2 have distinct functions and a complex consisting of FT and FDL1 mediates in photoperiodic control of seasonal growth. The downstream target of the FT-FD complex in photoperiodic control of growth is Like AP1 (LAP1), a tree ortholog of the floral meristem identity gene APETALA1. Intriguingly, FDL1 also participates in the transcriptional control of adaptive response and bud maturation pathways, independent of its interaction with FT, presumably via interaction with abscisic acid insensitive 3 (ABI3) transcription factor, a component of abscisic acid (ABA) signaling. Our data reveal that in contrast to its primary role in flowering, FD has dual roles in the photoperiodic control of seasonal growth and stress tolerance in trees. Thus, the functions of FT and FD have diversified during evolution, and FD homologs have acquired roles that are independent of their interaction with FT.

Estimation of the biomass and seasonal growth of external mycelium of ectomycorrhizal fungi in the field.

• In-growth mesh bags were used to quantify the production of external mycelium of ectomycorrhizal (EM) fungi in the field. • Colonization of the mesh bags was followed by visual estimation of the amount of mycelium, and by measuring fungal biomarkers (the phospholipid fatty acid (PLFA) 18 : 2ω6,9 and ergosterol). Mesh bags were placed inside and outside plots that were root isolated in order to estimate the amount of saprotrophic mycelium in relation to EM mycelium. The majority of mycelium in the mesh bags were EM, and this was confirmed by analysis of the δ13 C value in mycelia. • Fungal colonization of mesh bags peaked during autumn. The total amount of EM mycelium produced in the mesh bags during a year was calculated to be between 125 and 200 kg ha-1 . The total amount of EM mycelium (including EM mantles) in the humus was estimated to be 700-900 kg ha-1 . • The biomass of EM mycelium in the soil was in the same range as the biomass of fine roots and peaks of mycelial growth coincided with periods of maximum growth of fine-roots.

Meristem activity and biomass production as response mechanisms in two forest herbs.

Actaea spicata and Geranium sylvaticum are deciduous rosette herbs, frequently coexisting in nutrient-rich forests of boreal Sweden. A. spicata is restricted to this habitat whereas G. sylvaticum has a wider ecological amplitude. To explore the different distributions of these species we studied the growth mechanisms by which they responded to nutrient and light supply. Plants raised from seeds were grown under four combinations of light and nutrient supplies and growth was followed for 20 weeks. Growth responses were measured as biomass accumulation and meristem activity, i.e. leaf development by shoot meristems. Biomass accumulation and meristem activity were affected by nutrient and light treatments. However, A. spicata responded more to nutrients, whereas G. sylvaticum responded more to light, and low resource supply shortened the time during which shoot meristems of A. spicata developed leaves without a corresponding effect on biomass accumulation. In G. sylvaticum, the periods of leaf development and biomass accumulation were the same, but a relationship between high development rates and peak allocation of biomass to shoot meristems indicated a storage function of these during leaf development. We conclude that the inflexible growth of A. spicata, mainly affected by nutrient supply, makes it less competitive in open habitats and restricts it to shaded, nutrient-rich habitats. G. sylvaticum should be more successful in habitats subjected to unpredictable disturbances, due to a flexible growth morphology and a dynamic function of meristems that buffers against variation in nutrient supply and facilitates positive light responses. Meristematic data should be included in interspecific comparisons on growth responses to different resource supplies. Unless species have the same growth mechanism, it is not satisfactory to study biomass-related traits alone, as internal constraints concerning plant development might confuse the interpretation.

Nitrogen uptake, partitioning and internal cycling in Picea sitchensis (Bong.) Carr. as influenced by nitrogen supply.

Four-year-old seedlings of Picea sitchensis (Bong.) Carr. were grown in sand culture throughout 1989 and irrigated with nutrient solutions containing either 1.0 mol N m-3 (Low N) or 6.0 mol N m-3 (High N), to precondition their growth and capacity for N storage. During 1990 N enriched with 15 N was supplied, either from 15 March to 27 June, or 28 June to 20 November. Recovery of unlabelled N was used to determine the storage and immobilization of N for foliage growth, and the partitioning of labelled N taken up during the two periods was measured. Initial growth of trees in 1990 was unaffected by the current N supply and determined only by the N supplied the previous year. High N throughout increased the number of needles grown in 1990 compared to low N-treated trees, but had little effect on the dry weight of individual needles. When preconditioned with High N, trees responded to Low N in 1990 by a reduction in needle dry weight, without altering the number of needles produced. Low N trees supplied with High N in 1990 responded by increasing both needle numbers and dry weight, compared with trees supplied with Low N throughout. The amount of unlabelled N remobilized to foliage growth in 1990 was unaffected by the current N supply but reflected the amount of N in store, as determined by the N supply the previous year. The majority of N was remobilized from the 1989 foliage and none from roots. Partitioning of labelled N taken up during 1990 altered during the year, with a greater proportion of N taken up after 28 June recovered in the roots in all treatments, due to root growth as opposed to allocation of N to storage during the autumn, since root N concentrations fell between 17 June and the final harvest on 20 November.