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At least 199 records · Page 11

Phosphorus chemistry in plant charcoal: interplay between biomass composition and thermal condition

Background Vegetation fire may change Phosphorus (P) cycling in terrestrial ecosystems through converting biomass into fire residues. Aim The aim of this study was to understand the chemistry and mobility of P in fire residues to help reveal P thermochemistry during biomass burning and post-fire P cycling. Methods A combination of sequential extraction, liquid 31P NMR and P K-edge XANES was used to obtain quantitative P speciation and explain P solubilisation behaviours of charcoal. Key results Despite varying diverse P species existing in raw biomass, only two P structural moieties – orthophosphate and pyrophosphate – were identified in charcoal. However, relative abundance of pyrophosphate differs greatly among charcoal samples from different biomass types, ranging between 0 and 40% of total extractable P. Although P K-edge XANES data indicates abundant soluble phosphate minerals, most P (70–90%) is likely occluded physically in the charcoal. The bicarbonate-extractable P (the Olsen-P) varies significantly and cannot be explained by surface P concentration or elemental stoichiometry alone. Conclusion and implications The results suggest the importance of starting biomass P speciation (i.e. molecular structure and complexation environment) and thermal conditions in controlling P speciation and availability in charcoal. The different P chemistry between charcoal and ash suggests the importance of fire types and severity in disturbing the P cycle.

Forestry↗

Factors underlying a latitudinal gradient in the S/G lignin monomer ratio in natural poplar variants

The chemical composition of wood plays a pivotal role in the adaptability and structural integrity of trees. However, few studies have investigated the environmental factors that determine lignin composition and its biological significance in plants. Here, we examined the lignin syringyl-to-guaiacyl (S/G) ratio in members of a Populus trichocarpa population sourced from their native habitat and conducted a genome wide association study to identify genes linked to lignin formation. Our results revealed many significant associations, suggesting that lignin biosynthesis is a complex polygenic trait. Additionally, we found an increase in the S/G ratio from northern to southern geographic origin of the trees sampled, along with a corresponding metabolic and transcriptional reprogramming of xylem cell wall biosynthesis. Further molecular analysis identified a mutation in a cell wall laccase genetically associated with higher S/G ratios that predominate in trees from warmer lower latitudes. Collectively, our findings suggest that lignin heterogeneity arises from an evolutionary process enabling poplar adaptation to different climatic challenges.

adaptation↗

The Plant a Trillion Trees Campaign to Reduce Global Warming – Fleshing Out the Concept

The campaign to plant a trillion trees provided an easily understood approach to reduce the threat of global warming. However, focusing on trees does not consider that a maturing forest releases carbon (C) from dying trees offsetting C intake from growth of other trees, and results in only a one-time carbon storage benefit. Under sustainable management that harvests before forest growth slows down, forest C is transferred to wood products that substitute for fossil fuel (FF)- intensive products, reducing the carbon dioxide (CO 2 ) emissions with every harvest. Sustainably managed forests maintain growth by replanting (or regenerating), sustaining the C stored in products, as well as avoiding FF-intensive product emissions with every harvest. This paper addresses the question: Could planting more trees significantly reduce atmospheric CO 2 ; and if so, how? Survey data from many wood-producing mills show life cycle inventory (LCI) data on C that has been tracked through every stage of processing for many different wood products specific to each United States (USA) supply region, as well as C displacement measurements when substituting wood for FFintensive products. Results show that more and better-managed forest lands and more efficient product uses provide significant opportunities to reduce CO 2 emissions, especially in highly productive forest regions.

54 ENVIRONMENTAL SCIENCES↗

Seeing the forest for the trees: implementing dynamic representation of forest management and forest carbon in a long-term global multisector model

Abstract Studies have found that understanding forest management is critical in understanding the interaction between the carbon cycle and the integrated human-Earth system. This makes effectively representing forest management decisions such as planting and harvesting important. Here, we implement a novel dynamic forest harvest model in a global state of the art multi-sector dynamics model, namely the Global Change Analysis Model (GCAM). We implement an approach that explicitly tracks forest age and generates rotation ages for forest harvest that are responsive to changes in wood prices, changes in forest age and regional preferences for forest rotation. Furthermore, the forest sector in GCAM competes for investment with other land use types in the future years based on expected profit. Our baseline scenario results indicate that with the new forest harvest model, the current global wood product demand in GCAM can be met with minimal loss of old growth forest through the age-based harvest decisions. We find that economic pressure for deforestation and consequent loss of forest carbon is a bigger driver of global forest change than wood harvests, especially in developing regions. Under alternative scenarios where an economic value is placed on carbon across the terrestrial and energy systems, while there is an increase in forest plantations, there can be corresponding decreases in forest cover in some regions as forest land competes with land for bio-energy crops. When the carbon in forests is assigned a price, we find that the average rotation age for wood harvests can be reduced across regions to harvest forests in a more carbon efficient manner.

54 ENVIRONMENTAL SCIENCES↗

Thermal stress in degraded forests in the Brazilian Amazon Arc of Deforestation

Understanding thermal stress in tropical forests has taken on new urgency in light of accelerating climate change and expansion of deforestation and forest degradation. Degraded tropical forests in particular may be approaching critical temperature thresholds even more rapidly than intact forests, with implications for tree survival and ecosystem recovery. We investigate thermal stress in degraded tropical forests within the Brazilian Amazon Arc of Deforestation. Using land surface temperature data from the ECOsystem Spaceborne Thermal Radiometer Experiment on the international Space Station (ECOSTRESS), we compared canopy temperatures of intact, selectively logged, and burned forests in Feliz Natal, Mato Grosso, Brazil. Upper canopy temperatures in previously burned forests were 4.1% higher (mean = 36.5 °C) and 50.9% more variable compared to intact and logged forests, which showed remarkably similar temperature distributions (means of 34.9 °C and 35.1 °C, respectively). Modeled leaf temperature distributions based on canopy temperature measurements from one of the warmest days in a 2 year dry season record indicated that 87% of leaves in the warmest burned forest patches exceeded the temperature threshold where respiration surpasses photosynthesis, compared to approximately 72%–74% in intact and logged forests, respectively, in the same time period. After controlling for environmental factors, burned forests were predicted to be 2.6 °C warmer on average [1.39 °C–3.96 °C, 95% credible interval] than intact forests across a 5–40 m canopy height range. Burned forests showed modest thermal recovery over time, with temperatures decreasing by approximately 1.2 °C over a 30 year recovery period. In contrast, logged forests showed minimal thermal differences from intact forests (−0.353 on average, [−1.125–0.274, 95% credible interval]) and negligible change (0.15 °C) over the same timeframe. While the absolute probabilities of exceeding damaging thermal thresholds remain low across all forest types under current climate conditions, the probability of leaves exceeding temperatures that cause permanent leaf damage was ten times higher in burned forests, with implications for the future of burned forest regeneration in water-limited regions of the Amazon basin. In particular, the combination of higher mean temperatures, greater temperature variability, and more frequent exposure to damaging thermal thresholds implies that burned tropical forests will experience substantially higher mortality rates and slower biomass recovery compared to intact and selectively logged forests, especially in water-limited regions where trees cannot rely on evaporative cooling to moderate canopy temperatures.

Brazilian Amazon Arc of Deforestation↗

Deep-learning-based canopy height model generation from sub-meter resolution panchromatic satellite imagery

Canopy height models (CHMs) with sufficient resolution to distinguish individual trees are useful for a variety of applications. However, standard techniques to acquire such data, such as airborne lidar surveying, are often prohibitively expensive. Deep learning techniques for generating CHMs from high-resolution imagery are an attractive option to reduce costs. To date, success with these methods has been demonstrated using multichannel aerial photography and specialized satellite data products derived from multiple sensors, neither of which is commonly available at temporal resolutions finer than one year. Here we demonstrate a method to generate sub-meter resolution CHMs in three forests in California using a more abundant data source: sub-meter resolution, panchromatic satellite imagery from a single sensor. We show that phenology and species composition play important roles in model transferability; when trained using imagery from a single conifer forest in autumn, the model performs well on autumn imagery from a second conifer forest several hundred kilometers distant with no re-training. With modest additions to the training dataset, the same model generates minimally biased estimates of canopy height in both conifer and deciduous forests during multiple seasons. Because the model operates on satellite data with global coverage and a relatively short return interval, we propose its suitability to extrapolate tree-level canopy height data to remote regions and conduct high-temporal resolution monitoring of forest structure. We furthermore demonstrate the workflow’s applicability to fire modeling by conducting simulations in forests populated by trees measured using both this approach and airborne lidar surveying. We find minimal differences in fire behavior relative to a baseline case in which only statistical distributions of tree height and crown area are known. This result underscores the value of forest structural information derived from our workflow for improving the fidelity of wildland fire simulations, among other ecological applications.

54 ENVIRONMENTAL SCIENCES↗

Mentoring in the USDA Forest Service: A Survey of Aquatic Professionals

Abstract Mentoring is suggested as an important strategy to promote workplace inclusivity and is shown to be positively associated with high employee morale, yet mentee needs and experiences may not be universal. To evaluate mentoring impacts from the perspective of USDA Forest Service employees, we conducted an online survey of 251 aquatic professionals, including managers and scientists. 70% of respondents had mentors, and mentorship status did not vary across demographic characteristics. Previous mentoring relationships were most frequently identified as “informal” rather than “formal”; female employees were more likely to desire formal mentoring. Mentored respondents found their work more challenging, fulfilling, and valuable than unmentored respondents. Mentees looked for mentors who could provide constructive feedback, speak candidly, use active listening skills, and who cared about their careers. Overall, respondents were satisfied with their mentors’ skills. Despite strong demand for mentoring, access to mentors among aquatic professionals appears low across all categories.

Forestry↗

Long-term variability of root production in bioenergy crops from ingrowth core measurements

Long-term determination of root biomass production upon land-use conversion to biofuel crops is rare. To assess land-use legacy influences on belowground biomass accumulation, we converted 22-year-old Conservation Reserve Program (CRP) grasslands and 50+-year-old agricultural (AGR) lands to corn (C), switchgrass (Sw) and restored prairie (Pr) biofuel crops. We maintained one CRP grassland as a reference (Ref). We hypothesized that land-use history and crop type have significant effects on root density, with perennial crops on CRP grasslands having a higher root biomass productivity, while corn grown on former agricultural lands produce the lowest root biomass. Methods The ingrowth core method was used to determine in situ ingrowth root biomass, alongside measurements of aboveground net primary productivity (ANPP). Ancillary measurements, including air temperature, growing season length and precipitation were used to examine their influences on root biomass production. Important Findings Root biomass productivity was the highest in unconverted CRP grassland (1716 g m-2 yr-1) and lowest in corn fields (526 g m-2 yr-1). All perennial sites converted from CRP and AGR lands had lower root biomass and ANPP in the first year of planting but peaked in 2011 for switchgrass and a year later for restored prairies. Ecosystem stability was higher in restored prairies (AGR-Pr: 4.3 ± 0.11; CRP-Pr: 4.1 ± 0.10), with all monocultures exhibiting a lower stability. Root biomass production was positively related to ANPP (R2 = 0.40). Overall, attention should be given to root biomass accumulation in large-scale biofuel production as it is a major source of carbon sequestration.

Environmental Sciences & Ecology↗

Molecular mechanisms underlying host-induced gene silencing

Abstract Host-induced gene silencing (HIGS) refers to the silencing of genes in pathogens and pests by expressing homologous double-stranded RNAs (dsRNA) or artificial microRNAs (amiRNAs) in the host plant. The discovery of such trans-kingdom RNA silencing has enabled the development of RNA interference-based approaches for controlling diverse crop pathogens and pests. Although HIGS is a promising strategy, the mechanisms by which these regulatory RNAs translocate from plants to pathogens, and how they induce gene silencing in pathogens, are poorly understood. This lack of understanding has led to large variability in the efficacy of various HIGS treatments. This variability is likely due to multiple factors, such as the ability of the target pathogen or pest to take up and/or process RNA from the host, the specific genes and target sequences selected in the pathogen or pest for silencing, and where, when, and how the dsRNAs or amiRNAs are produced and translocated. In this review, we summarize what is currently known about the molecular mechanisms underlying HIGS, identify key unanswered questions, and explore strategies for improving the efficacy and reproducibility of HIGS treatments in the control of crop diseases.

59 BASIC BIOLOGICAL SCIENCES↗

Plants use molecular mechanisms mediated by biomolecular condensates to integrate environmental cues with development

This review highlights recent literature on biomolecular condensates in plant development and discusses challenges for fully dissecting their functional roles. Plant developmental biology has been inundated with descriptive examples of biomolecular condensate formation, but it is only recently that mechanistic understanding has been forthcoming. Here, we discuss recent examples of potential roles biomolecular condensates play at different stages of the plant life cycle. We group these examples based on putative molecular functions, including sequestering interacting components, enhancing dwell time, and interacting with cytoplasmic biophysical properties in response to environmental change. We explore how these mechanisms could modulate plant development in response to environmental inputs and discuss challenges and opportunities for further research into deciphering molecular mechanisms to better understand the diverse roles that biomolecular condensates exert on life.

59 BASIC BIOLOGICAL SCIENCES↗

A small molecule antagonizes jasmonic acid perception and auxin responses in vascular and nonvascular plants

The phytohormone jasmonoyl-L-isoleucine (JA-Ile) regulates many stress responses and developmental processes in plants. A co-receptor complex formed by the F-box protein Coronatine Insensitive 1 (COI1) and a Jasmonate (JA) ZIM-domain (JAZ) repressor perceives the hormone. JA-Ile antagonists are invaluable tools for exploring the role of JA-Ile in specific tissues and developmental stages, and for identifying regulatory processes of the signaling pathway. Using two complementary chemical screens, we identified three compounds that exhibit a robust inhibitory effect on both the hormone-mediated COI–JAZ interaction and degradation of JAZ1 and JAZ9 in vivo. One molecule, J4, also restrains specific JA-induced physiological responses in different angiosperm plants, including JA-mediated gene expression, growth inhibition, chlorophyll degradation, and anthocyanin accumulation. Interaction experiments with purified proteins indicate that J4 directly interferes with the formation of the Arabidopsis (Arabidopsis thaliana) COI1–JAZ complex otherwise induced by JA. The antagonistic effect of J4 on COI1–JAZ also occurs in the liverwort Marchantia polymorpha, suggesting the mode of action is conserved in land plants. Besides JA signaling, J4 works as an antagonist of the closely related auxin signaling pathway, preventing Transport Inhibitor Response1/Aux–indole-3-acetic acid interaction and auxin responses in planta, including hormone-mediated degradation of an auxin repressor, gene expression, and gravitropic response. However, J4 does not affect other hormonal pathways. Altogether, our results show that this dual antagonist competes with JA-Ile and auxin, preventing the formation of phylogenetically related receptor complexes. J4 may be a useful tool to dissect both the JA-Ile and auxin pathways in particular tissues and developmental stages since it reversibly inhibits these pathways. One-sentence summary: A chemical screen identified a molecule that antagonizes jasmonate perception by directly interfering with receptor complex formation in phylogenetically distant vascular and nonvascular plants.

59 BASIC BIOLOGICAL SCIENCES↗

Laisk measurements in the nonsteady state: Tests in plants exposed to warming and variable CO2 concentrations

Abstract Light respiration (RL) is an important component of plant carbon balance and a key parameter in photosynthesis models. RL is often measured using the Laisk method, a gas exchange technique that is traditionally employed under steady-state conditions. However, a nonsteady-state dynamic assimilation technique (DAT) may allow for more rapid Laisk measurements. In 2 studies, we examined the efficacy of DAT for estimating RL and the parameter Ci* (the intercellular CO2 concentration where Rubisco's oxygenation velocity is twice its carboxylation velocity), which is also derived from the Laisk technique. In the first study, we compared DAT and steady-state RL and Ci* estimates in paper birch (Betula papyrifera) growing under control and elevated temperature and CO2 concentrations. In the second, we compared DAT-estimated RL and Ci* in hybrid poplar (Populus nigra L. × P. maximowiczii A. Henry “NM6”) exposed to high or low CO2 concentration pre-treatments. The DAT and steady-state methods provided similar RL estimates in B. papyrifera, and we found little acclimation of RL to temperature or CO2; however, Ci* was higher when measured with DAT compared to steady-state methods. These Ci* differences were amplified by the high or low CO2 pre-treatments. We propose that changes in the export of glycine from photorespiration may explain these apparent differences in Ci*.

59 BASIC BIOLOGICAL SCIENCES↗

A century of studying plant secondary metabolism—From “what?” to “where, how, and why?”

Abstract Over the past century, early advances in understanding the identity of the chemicals that collectively form a living plant have led scientists to deeper investigations exploring where these molecules localize, how they are made, and why they are synthesized in the first place. Many small molecules are specific to the plant kingdom and have been termed plant secondary metabolites, despite the fact that they can play primary and essential roles in plant structure, development, and response to the environment. The past 100 yr have witnessed elucidation of the structure, function, localization, and biosynthesis of selected plant secondary metabolites. Nevertheless, many mysteries remain about the vast diversity of chemicals produced by plants and their roles in plant biology. From early work characterizing unpurified plant extracts, to modern integration of ‘omics technology to discover genes in metabolite biosynthesis and perception, research in plant (bio)chemistry has produced knowledge with substantial benefits for society, including human medicine and agricultural biotechnology. Here, we review the history of this work and offer suggestions for future areas of exploration. We also highlight some of the recently developed technologies that are leading to ongoing research advances.

54 ENVIRONMENTAL SCIENCES↗

Arabidopsis cytochrome b 5 proteins support fatty acid ω-3 but not ω-6 desaturation

Fatty acids are primary components of lipids, which serve as major energy sources in cells and play essential roles in membrane structure, signaling, and metabolic regulation (Shanklin and Cahoon 1998). The degree of fatty acid unsaturation critically influences lipid physicochemical properties, thereby affecting membrane fluidity and biological function (Nguyen et al. 2019). In Arabidopsis thaliana, fatty acid desaturation occurs via 2 parallel pathways: the “prokaryotic pathway” in plastids, involving glycosylglycerides, such as monogalactosyldiacylglycerol (MGDG) and digalactosyldiacylglycerol (DGDG), and phospholipid phosphatidylglycerol (PG); and the “eukaryotic pathway” in the endoplasmic reticulum (ER), involving phosphatidylcholine (PC) (Lou et al. 2014) (Supplementary Figure S1). Seven fatty acid desaturases (FADs) in Arabidopsis differentially desaturate each glycerolipid class in the plastid and ER (Nguyen et al. 2019). FAD2, an ER-resident ω-6 fatty acid desaturase, catalyzes the conversion of oleic acid (18:1) to linoleic acid (18:2), which can be further desaturated to α-linolenic acid (18:3) by FAD3, an ER-resident ω-3 fatty acid desaturase. In plastids, FAD6 catalyzes the desaturation of 18:1/16:1 to produce 18:2/16:2, while FAD7 and FAD8 redundantly convert 18:2/16:2 to 18:3/16:3 (Li-Beisson et al. 2013; Nguyen et al. 2019). Additionally, fatty acids synthesized in the ER can also be reimported into plastids to their site of de novo synthesis (Xu et al. 2010). All FADs require reducing power, in the form of 2 electrons, for catalysis, but the sources of the electrons vary between their subcellular localizations. In the ER, FAD2 and FAD3 receive electrons from a cytochrome b 5 (CB5)-based electron transfer chain comprising cytochrome b 5 reductase (CBR) and CB5. In contrast, ferredoxin serves as the electron donor for plastid-localized FAD6, FAD7, and FAD8 (Ohlrogge and Browse 1995; Andreu et al. 2007). While the relative contributions of the 2 pathways to total cellular desaturation products vary across tissues and species, most polyunsaturated FA biosynthesis in seeds occurs via ER-resident FAD2 and FAD3 (Miquel and Browse 1992; Ohlrogge and Browse 1995).

59 BASIC BIOLOGICAL SCIENCES↗

Short- and long-term responses of photosynthetic capacity to temperature in four boreal tree species in a free-air warming and rainfall manipulation experiment

Abstract High latitude forests cope with considerable variation in moisture and temperature at multiple temporal scales. To assess how their photosynthetic physiology responds to short- and long-term temperature variation, we measured photosynthetic capacity for four tree species growing in an open-air experiment in the boreal-temperate ecotone `Boreal Forest Warming at an Ecotone in Danger' (B4WarmED). The experiment factorially manipulated temperature above- and below-ground (ambient, +3.2 °C) and summer rainfall (ambient, 40% removal). We measured A/Ci curves at 18, 25 and 32 °C for individuals of two boreal (Pinus banksiana Lamb., Betula papyrifera Marsh.) and two temperate species (Pinus strobus L., Acer rubrum L.) experiencing the long-term warming and/or reduced-rainfall conditions induced by our experimental treatments. We calculated the apparent photosynthetic capacity descriptors VCmax,Ci and Jmax,Ci and their ratio for each measurement temperate. We hypothesized that (i) VCmax,Ci and Jmax,Ci would be down-regulated in plants experiencing longer term (e.g., weeks to months) warming and reduced rainfall (i.e., have lower values at a given measurement temperature), as is sometimes found in the literature, and that (ii) plants growing at warmer temperatures or from warmer ranges would show greater sensitivity (steeper slope) to short-term (minutes to hours) temperature variation. Neither hypothesis was supported as a general trend across the four species, as there was not a significant main effect (across species) of either warming or rainfall reduction on VCmax,Ci and Jmax,Ci. All species markedly increased VCmax,Ci and Jmax,Ci (and decreased their ratio) with short-term increases in temperature (i.e., contrasting values at 18, 25 and 32 °C), and those responses were independent of long-term treatments and did not differ among species. The Jmax,Ci:VCmax,Ci ratio was, however, significantly lower across species in warmed and reduced rainfall treatments. Collectively, these results suggest that boreal trees possess considerable short-term plasticity that may allow homeostasis of VCmax,Ci and Jmax,Ci to a longer term temperature treatment. Our results also caution against extrapolating results obtained under controlled and markedly contrasting temperature treatments to responses of photosynthetic parameters to more modest temperature changes expected in the near-term with climate warming in field conditions.

Forestry↗

Peak radial growth of diffuse-porous species occurs during periods of lower water availability than for ring-porous and coniferous trees

Climate models project warmer summer temperatures will increase the frequency and heat severity of droughts in temperate forests of Eastern North America. Hotter droughts are increasingly documented to affect tree growth and forest dynamics, with critical impacts on tree mortality, carbon sequestration and timber provision. The growing acknowledgement of the dominant role of drought timing on tree vulnerability to water deficit raises the issue of our limited understanding of radial growth phenology for most temperate tree species. Here, we use well-replicated dendrometer band data sampled frequently during the growing season to assess the growth phenology of 610 trees from 15 temperate species over 6 years. Patterns of diameter growth follow a typical logistic shape, with growth rates reaching a maximum in June, and then decreasing until process termination. On average, we find that diffuse-porous species take 16–18 days less than other wood-structure types to put on 50% of their annual diameter growth. However, their peak growth rate occurs almost a full month later than ring-porous and conifer species (ca. 24 ± 4 days; mean ± 95% credible interval). Unlike other species, the growth phenology of diffuse-porous species in our dataset is highly correlated with their spring foliar phenology. We also find that the later window of growth in diffuse-porous species, coinciding with peak evapotranspiration and lower water availability, exposes them to a higher water deficit of 88 ± 19 mm (mean ± SE) during their peak growth than ring-porous and coniferous species (15 ± 35 mm and 30 ± 30 mm, respectively). Given the high climatic sensitivity of wood formation, our findings highlight the importance of wood porosity as one predictor of species climatic sensitivity to the projected intensification of the drought regime in the coming decades.

59 BASIC BIOLOGICAL SCIENCES↗

Modeling starch dynamics from seasonal variations of photosynthesis, growth and respiration

Abstract Nonstructural carbohydrates (NSCs) buffer differences in plant carbon supply (photosynthesis) and demand (respiration, growth, etc.), but the regulation of their dynamics remains unresolved. Seasonal variations in NSCs are well-documented, but differences in the time-average, amplitude, phase and other characteristics across ecosystems and functional types lack explanation; furthermore, observed dynamics do not always match expectations. The failure to match observed and expected dynamics has stimulated debate on whether carbon supply or demand drives NSC dynamics. To gain insight into how carbon supply and demand drive seasonal NSC dynamics, we derive a simple model of NSC dynamics based on carbon mass balance and linearizing the NSC demand to determine how supply-driven and demand-driven seasonal NSC dynamics differ. We find that supply-driven and demand-driven dynamics yield distinct timings of seasonal extrema, and supply overrides demand when carbon supply is low in winter (e.g., at high latitudes). Our results also suggest that NSC dynamics often lag changes carbon mass balance. We also predict differences in NSC dynamics across mass, suggesting that saplings are more dynamic and respond more quickly to the environment than mature trees. Our findings suggest that substrate-dependent regulation with environmental variation is sufficient to generate complex NSC dynamics.

Forestry↗