A unified explanation for the morphology of raised peatlands.

Raised peatlands, or bogs, are gently mounded landforms that are composed entirely of organic matter1-4 and store the most carbon per area of any terrestrial ecosystem5. The shapes of bogs are critically important because their domed morphology4,6,7 accounts for much of the carbon that bogs store and determines how they will respond to interventions8,9 to stop greenhouse gas emissions and fires after anthropogenic drainage10-13. However, a general theory to infer the morphology of bogs is still lacking4,6,7. Here we show that an equation based on the processes universal to bogs explains their morphology across biomes, from Alaska, through the tropics, to New Zealand. In contrast to earlier models of bog morphology that attempted to describe only long-term equilibrium shapes4,6,7 and were, therefore, inapplicable to most bogs14-16, our approach makes no such assumption and makes it possible to infer full shapes of bogs from a sample of elevations, such as a single elevation transect. Our findings provide a foundation for quantitative inference about the morphology, hydrology and carbon storage of bogs through Earth's history, as well as a basis for planning natural climate solutions by rewetting damaged bogs around the world.

Genetic structure and population history of a peat swamp forest tree species, Shorea albida (Dipterocarpaceae), in Brunei Darussalam.

Southeast Asia supports high biodiversity, in a mosaic of forest types formed by the expansion and contraction of habitats through past climate changes. Among the region's forest types, the geographical distribution of peat swamp forests has fluctuated intensely over the past 120,000 years. Most peat swamp forests in Southeast Asia are found in coastal regions and formed within the last 7,000 years after a decline in sea level. However, some peat swamps were initiated earlier on substrates of slightly higher elevation, and these peat swamps might have been refugia for peat swamp species in the last glacial period and the high sea level period. We assessed genetic diversity, genetic structure and divergence time of current genetic groups for Shorea albida in Brunei, an endemic tree species of Bornean peat swamp forests, using 18 microsatellite markers. Genetic diversity was not lower than has been found in other Shorea species, possibly because of the high density of S. albida in Brunei. Although overall genetic divergence between populations was low, two populations (Ingei and Labi Road 3) were distinct from the other populations. Analysis using DIYABC estimated that three genetic groups (Ingei, Labi Road 3 and others) diverged simultaneously from their ancestral population, whose effective size was very small, about 7,500 years ago, corresponding to a recent sea level peak in the Belait-Baram river basin. In that high sea level period, some higher-elevation lands remained, and peat formation had already started in this region. We propose that the current genetic structure of S. albida in Brunei was formed from small refugial populations that survived the period of higher sea level in these higher-elevation areas. Because of their relatively high genetic diversity, Brunei's S. albida populations should become an important genetic resource for the recovery of genetically healthy populations in other parts of northwest Borneo.

Net greenhouse gas balance of fibre wood plantation on peat in Indonesia.

Tropical peatlands cycle and store large amounts of carbon in their soil and biomass1-5. Climate and land-use change alters greenhouse gas (GHG) fluxes of tropical peatlands, but the magnitude of these changes remains highly uncertain6-19. Here we measure net ecosystem exchanges of carbon dioxide, methane and soil nitrous oxide fluxes between October 2016 and May 2022 from Acacia crassicarpa plantation, degraded forest and intact forest within the same peat landscape, representing land-cover-change trajectories in Sumatra, Indonesia. This allows us to present a full plantation rotation GHG flux balance in a fibre wood plantation on peatland. We find that the Acacia plantation has lower GHG emissions than the degraded site with a similar average groundwater level (GWL), despite more intensive land use. The GHG emissions from the Acacia plantation over a full plantation rotation (35.2 ± 4.7 tCO2-eq ha-1 year-1, average ± standard deviation) were around two times higher than those from the intact forest (20.3 ± 3.7 tCO2-eq ha-1 year-1), but only half of the current Intergovernmental Panel on Climate Change (IPCC) Tier 1 emission factor (EF)20 for this land use. Our results can help to reduce the uncertainty in GHG emissions estimates, provide an estimate of the impact of land-use change on tropical peat and develop science-based peatland management practices as nature-based climate solutions.

Variation in carbon and nitrogen concentrations among peatland categories at the global scale.

Peatlands account for 15 to 30% of the world's soil carbon (C) stock and are important controls over global nitrogen (N) cycles. However, C and N concentrations are known to vary among peatlands contributing to the uncertainty of global C inventories, but there are few global studies that relate peatland classification to peat chemistry. We analyzed 436 peat cores sampled in 24 countries across six continents and measured C, N, and organic matter (OM) content at three depths down to 70 cm. Sites were distinguished between northern (387) and tropical (49) peatlands and assigned to one of six distinct broadly recognized peatland categories that vary primarily along a pH gradient. Peat C and N concentrations, OM content, and C:N ratios differed significantly among peatland categories, but few differences in chemistry with depth were found within each category. Across all peatlands C and N concentrations in the 10-20 cm layer, were 440 ± 85.1 g kg-1 and 13.9 ± 7.4 g kg-1, with an average C:N ratio of 30.1 ± 20.8. Among peatland categories, median C concentrations were highest in bogs, poor fens and tropical swamps (446-532 g kg-1) and lowest in intermediate and extremely rich fens (375-414 g kg-1). The C:OM ratio in peat was similar across most peatland categories, except in deeper samples from ombrotrophic tropical peat swamps that were higher than other peatlands categories. Peat N concentrations and C:N ratios varied approximately two-fold among peatland categories and N concentrations tended to be higher (and C:N lower) in intermediate fens compared with other peatland types. This study reports on a unique data set and demonstrates that differences in peat C and OM concentrations among broadly classified peatland categories are predictable, which can aid future studies that use land cover assessments to refine global peatland C and N stocks.

CO2 emissions from an undrained tropical peatland: Interacting influences of temperature, shading and water table depth.

Emission of CO2 from tropical peatlands is an important component of the global carbon budget. Over days to months, these fluxes are largely controlled by water table depth. However, the diurnal cycle is less well understood, in part, because most measurements have been collected daily at midday. We used an automated chamber system to make hourly measurements of peat surface CO2 emissions from chambers root-cut to 30 cm. We then used these data to disentangle the relationship between temperature, water table and heterotrophic respiration (Rhet ). We made two central observations. First, we found strong diurnal cycles in CO2 flux and near-surface peat temperature (<10 cm depth), both peaking at midday. The magnitude of diurnal oscillations was strongly influenced by shading and water table depth, highlighting the limitations of relying on daytime measurements and/or a single correction factor to remove daytime bias in flux measurements. Second, we found mean daily Rhet had a strong linear relationship to the depth of the water table, and under flooded conditions, Rhet was small and constant. We used this relationship between Rhet and water table depth to estimate carbon export from both Rhet and dissolved organic carbon over the course of a year based on water table records. Rhet dominates annual carbon export, demonstrating the potential for peatland drainage to increase regional CO2 emissions. Finally, we discuss an apparent incompatibility between hourly and daily average observations of CO2 flux, water table and temperature: water table and daily average flux data suggest that CO2 is produced across the entire unsaturated peat profile, whereas temperature and hourly flux data appear to suggest that CO2 fluxes are controlled by very near surface peat. We explore how temperature-, moisture- and gas transport-related mechanisms could cause mean CO2 emissions to increase linearly with water table depth and also have a large diurnal cycle.

Tropical peatland carbon storage linked to global latitudinal trends in peat recalcitrance.

Peatlands represent large terrestrial carbon banks. Given that most peat accumulates in boreal regions, where low temperatures and water saturation preserve organic matter, the existence of peat in (sub)tropical regions remains enigmatic. Here we examined peat and plant chemistry across a latitudinal transect from the Arctic to the tropics. Near-surface low-latitude peat has lower carbohydrate and greater aromatic content than near-surface high-latitude peat, creating a reduced oxidation state and resulting recalcitrance. This recalcitrance allows peat to persist in the (sub)tropics despite warm temperatures. Because we observed similar declines in carbohydrate content with depth in high-latitude peat, our data explain recent field-scale deep peat warming experiments in which catotelm (deeper) peat remained stable despite temperature increases up to 9 °C. We suggest that high-latitude deep peat reservoirs may be stabilized in the face of climate change by their ultimately lower carbohydrate and higher aromatic composition, similar to tropical peats.

Maximum-likelihood estimation of xylem vessel length distributions.

Vessel length is an important functional trait for plant hydraulics, because it determines the ratio of flow resistances posed by lumen and pit membranes and hence controls xylem hydraulic efficiency. The most commonly applied methods to estimate vessel lengths are based on the injection of silicon or paint into cut-off stem segments. The number of stained vessels in a series of cross-sections in increasing distance from the injection point is then counted. The resulting infusion profiles are used to estimate the vessel length distribution using one of several statistical algorithms. However, the basis of these algorithms has not been systematically analysed using probability theory. We derive a general mathematical expression for the expected shape of the infusion profile for a given vessel length distribution, provide analytic solutions for five candidate distributions (exponential, Erlang(2), gamma, Weibull, and log-normal), and present maximum likelihood estimators for the parameters of these distributions including implementations in R based on two potential sampling schemes (counting all injected vessels or counting the injected and empty vessels in a random subset of each cross-section). We then explore the performance of these estimators relative to other methods with Monte Carlo experiments. Our analysis demonstrates that most published methods estimate the conditional length distribution of vessels that cross an injection point, which is a size-biased version of the overall length distribution in the stem. We show the mathematical relationship between these distributions and provide methods to estimate either of them. According to our simulation experiments, vessel length distribution was best described by the more flexible models, especially the Weibull distribution. In simulations, the estimators were able to recover the parameters of the vessel length distribution if its functional form was known, achieving an overlap of 90% or more between the true and predicted length distribution when counting no more than 500 injected vessels in 10 cross-sections. This sample size nowadays can easily be reached with the help of automated image analysis.

How temporal patterns in rainfall determine the geomorphology and carbon fluxes of tropical peatlands.

Tropical peatlands now emit hundreds of megatons of carbon dioxide per year because of human disruption of the feedbacks that link peat accumulation and groundwater hydrology. However, no quantitative theory has existed for how patterns of carbon storage and release accompanying growth and subsidence of tropical peatlands are affected by climate and disturbance. Using comprehensive data from a pristine peatland in Brunei Darussalam, we show how rainfall and groundwater flow determine a shape parameter (the Laplacian of the peat surface elevation) that specifies, under a given rainfall regime, the ultimate, stable morphology, and hence carbon storage, of a tropical peatland within a network of rivers or canals. We find that peatlands reach their ultimate shape first at the edges of peat domes where they are bounded by rivers, so that the rate of carbon uptake accompanying their growth is proportional to the area of the still-growing dome interior. We use this model to study how tropical peatland carbon storage and fluxes are controlled by changes in climate, sea level, and drainage networks. We find that fluctuations in net precipitation on timescales from hours to years can reduce long-term peat accumulation. Our mathematical and numerical models can be used to predict long-term effects of changes in temporal rainfall patterns and drainage networks on tropical peatland geomorphology and carbon storage.

Tensioning the helix: a mechanism for force generation in twining plants.

Twining plants use their helical stems to clasp supports and to generate a squeezing force, providing stability against gravity. To elucidate the mechanism that allows force generation, we measured the squeezing forces exerted by the twiner Dioscorea bulbifera while following its growth using time-lapse photography. We show that the development of the squeezing force is accompanied by stiffening of the stem and the expansion of stipules at the leaf base. We use a simple thin rod model to show that despite their small size and sparse distribution, stipules impose a stem deformation sufficient to account for the measured squeezing force. We further demonstrate that tensioning of the stem helix, although counter-intuitive, is the most effective mechanism for generating large squeezing forces in twining plants. Our observations and model point to a general mechanism for the generation of the twining force: a modest radial stem expansion during primary growth, or the growth of lateral structures such as leaf bases, causes a delayed stem tensioning that creates the squeezing forces necessary for twining plants to ascend their supports. Our study thus provides the long-sought answer to the question of how twining plants ascend smooth supports without the use of adhesive or hook-like structures.

Structure and function of bordered pits: new discoveries and impacts on whole-plant hydraulic function.

Bordered pits are cavities in the lignified cell walls of xylem conduits (vessels and tracheids) that are essential components in the water-transport system of higher plants. The pit membrane, which lies in the center of each pit, allows water to pass between xylem conduits but limits the spread of embolism and vascular pathogens in the xylem. Averaged across a wide range of species, pits account for > 50% of total xylem hydraulic resistance, indicating that they are an important factor in the overall hydraulic efficiency of plants. The structure of pits varies dramatically across species, with large differences evident in the porosity and thickness of pit membranes. Because greater porosity reduces hydraulic resistance but increases vulnerability to embolism, differences in pit structure are expected to correlate with trade-offs between efficiency and safety of water transport. However, trade-offs in hydraulic function are influenced both by pit-level differences in structure (e.g. average porosity of pit membranes) and by tissue-level changes in conduit allometry (average length, diameter) and the total surface area of pit membranes that connects vessels. In this review we address the impact of variation in pit structure on water transport in plants from the level of individual pits to the whole plant.

Dynamics of freeze-thaw embolism in Smilax rotundifolia (Smilacaceae).

Freeze-thaw cycles pose a major physiological challenge for all temperate perennial plants, but monocotyledonous vines face a still greater risk because their few large vessels are especially susceptible to embolism and are not replaced by secondary growth. The genus Smilax is particularly remarkable because it is widespread in the tropics but includes species that survive the hard frosts of New England winters. Smilax rotundifolia was monitored for a year for evidence of stem xylem freeze-thaw cavitation and refilling. Embolism of metaxylem was complete by late November and was completely reversed by late April, when root pressures rose as high as 100 kPa. Protoxylem remained full of sap throughout the year in cryogenic scanning electron micrographs. Three methods were used to quantify embolism: percent loss conductivity (PLC), gravimetric air fraction (GAF: mass of water in stem xylem relative to capacity), and cryogenic scanning electron microscopy (cryo-SEM). The three methods corroborated one another well and gave quantitatively similar results. Osmolality of xylem sap extracted from exuding stems was 64 mol/kg (±7.0, N = 8), consistent with the root pressures observed. Strong root pressure can account for Smilax's survival in temperate regions with severe frosts, where few monocots with persistent aboveground organs are found.

Direct measurements of intervessel pit membrane hydraulic resistance in two angiosperm tree species.

The hydraulic resistance of pit membranes was measured directly in earlywood vessels of Fraxinus americana and Ulmus americana. The area-specific resistance of pit membranes (r(mem)) was higher than modeled or measured values obtained previously for hardwood species, with r(mem) of 5.24 × 10(3) MPa·s·m(-1) for Fraxinus and 2.56 × 10(3) MPa·s·m(-1) for Ulmus. The calculated resistance of pit canals was three orders of magnitude below total pit resistance indicating that pit membranes contributed the majority of resistance. Scanning electron microscopy indicated that pit membranes of Ulmus were thinner and more porous than those of Fraxinus, consistent with the difference in r(mem) between the species. Measurements of average vessel diameter and length and area of wall overlap with neighboring vessels were used to partition the vascular resistance between vessel lumen and pit membrane components. Pit membrane resistance accounted for 80% of the total resistance in Fraxinus and 87% in Ulmus in 2-yr-old branch sections. However, measurements of vessel dimensions in the trunk suggest that the division of resistance between pit membrane and lumen components would be closer to co-limiting in older regions of the tree. Thus, pit membrane resistance may be of greater relative importance in small branches than in older regions of mature trees.