Phylogenomic structure and speciation in an emerging model: the Sphagnum magellanicum complex (Bryophyta).

Sphagnum magellanicum is one of two Sphagnum species for which a reference-quality genome exists to facilitate research in ecological genomics. Phylogenetic and comparative genomic analyses were conducted based on resequencing data from 48 samples and RADseq analyses based on 187 samples. We report herein that there are four clades/species within the S. magellanicum complex in eastern North America and that the reference genome belongs to Sphagnum divinum. The species exhibit tens of thousands (RADseq) to millions (resequencing) of fixed nucleotide differences. Two species, however, referred to informally as S. diabolicum and S. magni because they have not been formally described, are differentiated by only 100 (RADseq) to 1000 (resequencing) of differences. Introgression among species in the complex is demonstrated using D-statistics and f4 ratios. One ecologically important functional trait, tissue decomposability, which underlies peat (carbon) accumulation, does not differ between segregates in the S. magellanicum complex, although previous research showed that many closely related Sphagnum species have evolved differences in decomposability/carbon sequestration. Phylogenetic resolution and more accurate species delimitation in the S. magellanicum complex substantially increase the value of this group for studying the early evolutionary stages of climate adaptation and ecological evolution more broadly.

Ecophysiology of leaf trichomes.

This review examines how leaf trichomes influence leaf physiological responses to abiotic environmental drivers. Leaf trichomes are known to modulate leaf traits, particularly radiation absorptance, but studies in recent decades have demonstrated that trichomes have a more expansive role in the plant-environment interaction. Although best known as light reflectors, dense trichome canopies modulate leaf heat balance and photon interception, and consequently affect gas exchange traits. Analysis of published studies shows that dense pubescence generally increases reflectance of visible light and near-infrared and infrared radiation. Reflective trichomes are also protective, reducing photoinhibition and UV-B related damage to leaf photochemistry. Little support exists for a strong trichome effect on leaf boundary layer resistance and transpiration, but recent studies indicate they may play a substantive role in leaf water relations affecting leaf wettability, droplet retention and leaf water uptake. Different lines of evidence indicate that adaxial and abaxial trichomes may function quite differently, even within the same leaf. Overall, this review synthesises and re-examines the diverse array of relevant studies from the past 40 years, illustrating our current understanding of how trichomes influence the energy, carbon and water balance of plants, and highlighting promising areas for future research.

Regulation and acclimation of leaf gas exchange in a piñon-juniper woodland exposed to three different precipitation regimes.

Leaf gas-exchange regulation plays a central role in the ability of trees to survive drought, but forecasting the future response of gas exchange to prolonged drought is hampered by our lack of knowledge regarding potential acclimation. To investigate whether leaf gas-exchange rates and sensitivity to drought acclimate to precipitation regimes, we measured the seasonal variations of leaf gas exchange in a mature piñon-juniper Pinus edulis-Juniperus monosperma woodland after 3 years of precipitation manipulation. We compared trees receiving ambient precipitation with those in an irrigated treatment (+30% of ambient precipitation) and a partial rainfall exclusion (-45%). Treatments significantly affected leaf water potential, stomatal conductance and photosynthesis for both isohydric piñon and anisohydric juniper. Leaf gas exchange acclimated to the precipitation regimes in both species. Maximum gas-exchange rates under well-watered conditions, leaf-specific hydraulic conductance and leaf water potential at zero photosynthetic assimilation all decreased with decreasing precipitation. Despite their distinct drought resistance and stomatal regulation strategies, both species experienced hydraulic limitation on leaf gas exchange when precipitation decreased, leading to an intraspecific trade-off between maximum photosynthetic assimilation and resistance of photosynthesis to drought. This response will be most detrimental to the carbon balance of piñon under predicted increases in aridity in the southwestern USA.

δ(13) C of leaf-respired CO(2) reflects intrinsic water-use efficiency in barley.

Leaf intrinsic water-use efficiency (WUE), the ratio of photosynthetic rate to stomatal conductance (A/g(s) ), is a key plant trait linking terrestrial carbon and water cycles. A rapid, integrative proxy for A/g(s) is of benefit to crop breeding programmes aiming to improve WUE, but also for ecologists interested in plant carbon-water balance in natural systems. We hypothesize that the carbon isotope composition of leaf-respired CO(2) (δ(13) C(Rl) ), two hours after leaves are transferred to the dark, records photosynthetic carbon isotope discrimination and so provides a proxy for A/g(s) . To test this hypothesis, δ(13) C(Rl) was measured in four barley cultivars grown in the field at two levels of water availability and compared to leaf-level gas exchange (the ratio of leaf intercellular to ambient CO(2) partial pressure, C(i) /C(a) , and A/g(s) ). Leaf-respired CO(2) was more (13) C-depleted in plants grown at higher water availability, varied between days as environmental conditions changed, and was significantly different between cultivars. A strong relationship between δ(13) C(Rl) and δ(13) C of sucrose was observed. δ(13) C(Rl) was converted into apparent photosynthetic discrimination (Δ(13) C(Rl) ) revealing strong relationships between Δ(13) C(Rl) and C(i) /C(a) and A/g(s) during the vegetative stage of growth. We therefore conclude that δ(13) C(Rl) may provide a rapid, integrative proxy for A/g(s) in barley.

Influence of diurnal variation in mesophyll conductance on modelled 13C discrimination: results from a field study.

Mesophyll conductance to CO(2) (g(m)) limits carbon assimilation and influences carbon isotope discrimination (Delta) under most environmental conditions. Current work is elucidating the environmental regulation of g(m), but the influence of g(m) on model predictions of Delta remains poorly understood. In this study, field measurements of Delta and g(m) were obtained using a tunable diode laser spectroscope coupled to portable photosynthesis systems. These data were used to test the importance of g(m) in predicting Delta using the comprehensive Farquhar model of Delta (Delta(comp)), where g(m) was parameterized using three methods based on: (i) mean g(m); (ii) the relationship between stomatal conductance (g(s)) and g(m); and (iii) the relationship between time of day (TOD) and g(m). Incorporating mean g(m), g(s)-based g(m), and TOD-based g(m) did not consistently improve Delta(comp) predictions of field-grown juniper compared with the simple model of Delta (Delta(simple)) that omits fractionation factors associated with g(m) and decarboxylation. Sensitivity tests suggest that b, the fractionation due to carboxylation, was lower (25 per thousand) than the value commonly used in Delta(comp) (29 per thousand) and Delta(simple) (27 per thousand). These results demonstrate the limits of all tested models in predicting observed juniper Delta, largely due to unexplained offsets between predicted and observed values that were not reconciled in sensitivity tests of variability in g(m), b, or e, the day respiratory fractionation.

High-frequency field measurements of diurnal carbon isotope discrimination and internal conductance in a semi-arid species, Juniperus monosperma.

We present field observations of carbon isotope discrimination (Delta) and internal conductance of CO(2) (g(i)) collected using tunable diode laser spectroscopy (TDL). Delta ranged from 12.0 to 27.4 per thousand over diurnal periods with daily means from 16.3 +/- 0.2 per thousand during drought to 19.0 +/- 0.5 per thousand during monsoon conditions. We observed a large range in g(i), with most estimates between 0.04 and 4.0 micromol m(-2) s(-1) Pa(-1). We tested the comprehensive Farquhar, O'Leary and Berry model of Delta (Delta(comp)), a simplified form of Delta(comp) (Delta(simple)) and a recently suggested amendment (Delta(revised)). Sensitivity analyses demonstrated that varying g(i) had a substantial effect on Delta(comp), resulting in mean differences between observed Delta (Delta(obs)) and Delta(comp) ranging from 0.04 to 9.6 per thousand. First-order regressions adequately described the relationship between Delta and the ratio of substomatal to atmospheric CO(2) partial pressure (p(i)/p(a)) on all 3 d, but second-order models better described the relationship in July and August. The three tested models each best predicted Delta(obs) on different days. In June, Delta(simple) outperformed Delta(comp) and Delta(revised), but incorporating g(i) and all non-photosynthetic fractionations improved model predictions in July and August.

A new measurement technique reveals temporal variation in delta18O of leaf-respired CO2.

The oxygen isotope composition of CO(2) respired by Ricinus communis leaves (delta(18)O(R)) was measured under non-steady-state conditions with a temporal resolution of 3 min using a tunable diode laser (TDL) absorption spectrometer coupled to a portable gas exchange system. The SD of delta(18)O measurement by the TDL was +/- 0.2 per thousand and close to that of traditional mass spectrometers. Further, delta(18)O(R) values at isotopic steady state were comparable to those obtained using traditional flask sampling and mass spectrometric techniques for R. communis grown and measured in similar environmental conditions. As well as higher temporal resolution, the online TDL method described here has a number of advantages over mass spectrometric techniques. At isotopic steady state among plants grown at high light, the "one-way flux" model was required to accurately predict delta(18)O(R). A comparison of measurements and the model suggests that plants grown under low-light conditions have either a lower proportion of chloroplast CO(2) that isotopically equilibrates with chloroplast water, or more enriched delta(18)O of CO(2) in the chloroplast that has not equilibrated with local water. The high temporal resolution of isotopic measurements allowed the first measurements of delta(18)O(R) when stomatal conductance was rapidly changing. Under non-steady-state conditions, delta(18)O(R) varied between 50 and 220 per thousand for leaves of plants grown under different light and water environments, and varied by as much as 100 per thousand within 10 min for a single leaf. Stomatal conductance ranged from 0.001 to 1.586 mol m(-2) s(-1), and had an important influence on delta(18)O(R) under non-steady-state conditions not only via effects on leaf water H(2) (18)O enrichment, but also via effects on the rate of the one-way fluxes of CO(2) into and out of the leaf.

A new measurement technique reveals rapid post-illumination changes in the carbon isotope composition of leaf-respired CO2.

We describe an open leaf gas exchange system coupled to a tunable diode laser (TDL) spectroscopy system enabling measurement of the leaf respiratory CO(2) flux and its associated carbon isotope composition (delta(13)C(Rl)) every 3 min. The precision of delta(13)C(Rl) measurement is comparable to that of traditional mass spectrometry techniques. delta(13)C(Rl) from castor bean (Ricinus communis L.) leaves tended to be positively related to the ratio of CO(2) produced to O(2) consumed [respiratory quotient (RQ)] after 24-48 h of prolonged darkness, in support of existing models. Further, the apparent fractionation between respiratory substrates and respired CO(2) within 1-8 h after the start of the dark period was similar to previous observations. In subsequent experiments, R. communis plants were grown under variable water availability to provide a range in delta(13)C of recently fixed carbohydrate. In leaves exposed to high light levels prior to the start of the dark period, CO(2) respired by leaves was up to 11 per thousand more enriched than phloem sap sugars within the first 10-15 min after plants had been moved from the light into the dark. The (13)C enrichment in respired CO(2) then decreased rapidly to within 3-7 per thousand of phloem sap after 30-60 min in the dark. This strong enrichment was not observed if light levels were low prior to the start of the dark period. Measurements of RQ confirmed that carbohydrates were the likely respiratory substrate for plants (RQ > 0.8) within the first 60 min after illumination. The strong (13)C enrichment that followed a high light-to-dark transition coincided with high respiration rates, suggesting that so-called light-enhanced dark respiration (LEDR) is fed by (13)C-enriched metabolites.

Host physiological condition regulates parasitic plant performance: Arceuthobium vaginatum subsp. cryptopodum on Pinus ponderosa.

Much research has focused on effects of plant parasites on host-plant physiology and growth, but little is known about effects of host physiological condition on parasite growth. Using the parasitic dwarf mistletoe Arceuthobium vaginatum subsp. cryptopodum (Viscaceae) and its host Pinus ponderosa, we investigated whether changes in host physiological condition influenced mistletoe shoot development in northern Arizona forests. We conducted two studies in two consecutive years and used forest thinning (i.e., competitive release) to manipulate host physiological condition. We removed dwarf mistletoe shoots in April, before the onset of the growing season, and measured the amount of regrowth in the first season after forest thinning (Study I: n=38 trees; Study II: n=35 trees). Thinning increased tree uptake of water and carbon in both studies, but had no effect on leaf N concentration or delta13C. Mistletoe shoot growth was greater on trees with high uptake of water and carbon in thinned stands than trees with low uptake in unthinned stands. These findings show that increased resource uptake by host trees increases resources to these heterotrophic dwarf mistletoes, and links mistletoe performance to changes in host physiological condition.