Evolutionary Physiology: The extent of C4 and CAM photosynthesis in the Genera Anacampseros and Grahamia of the Portulacaceae

The Portulacaceae is one of the few terrestrial plant families known to have both C(4) and Crassulacean acid metabolism (CAM) species. There may be multiple origins of the evolution of CAM within the Portulacaceae but the only clear evidence of C(4) photosynthesis is found in members of the genus Portulaca. In the Portulaca, CAM succulent tissue is overlaid with the C(4) tissue in a unique fashion where both pathways are operating simultaneously. Earlier reports have shown that the clade containing the genera Anacampseros and Grahamia may also contain C(4) photosynthetic species similar to the Portulaca, which would indicate multiple origins of C(4) photosynthesis within the family. The aim of the present study was to ascertain the true photosynthetic nature of these genera. An initial survey of the carbon isotope composition of the Anacampseros ranged from -12.6 per thousand to -24.0 per thousand, indicating very little CAM activity in some species, with other values close to the C(4) range. Anacampseros (=Grahamia) australiana which had been previously identified as a C(4) species had a carbon isotope composition value of -24.0 per thousand, which is more indicative of a C(3) species with a slight contribution of CAM activity. Other Anacampseros species with C(4)-like values have been shown to be CAM plants. The initial isotope analysis of the Grahamia species gave values in the range of -27.1 per thousand to -23.6 per thousand, placing the Grahamia species well towards the C(3) photosynthetic range. Further physiological studies indicated increased night-time CO(2) uptake with imposition of water stress, associated with a large diurnal acid fluctuation and a marked increased phosphoenolpyruvate carboxylase activity. This showed that the Grahamia species are actually facultative CAM plants despite their C(3)-like carbon isotope values. The results indicate that the Grahamia and Anacampseros species do not utilize the C(4) photosynthetic pathway. This is the first to identify that the Grahamia species are facultative CAM plants where CAM can be induced by water stress. This work supports earlier physiological work that indicates that this clade containing Anacampseros and Grahamia species comprises predominantly facultative CAM plants. This report suggests there may be only one clade which contains C(4) photosynthetic members with CAM-like characteristics

The sensitivity of photosynthesis to O2 and CO2 concentration identifies strong Rubisco control above the thermal optimum

This work was funded by an NSERC Discovery and an NSERC Accelerator grant to R.F.S. F.A.B. is supported by the ARC Centre of Excellence for Translational Photosynthesis

The effect of carbon and nutrient loading during nursery culture on the growth of black spruce seedlings: a six-year field study

Abstract We tested the effects of exponential nutrient loading and springtime carbon loading during nursery culture on the field performance of black spruce (Picea mariana (Mill.) B.S.P.). Seedlings were grown from seed with a conventional, fixed dose fertilizer (10 mg N seedling À1 ) or an exponential nutrient loading regime (75 mg N seedling À1 ). The following spring, seedlings were exposed for two weeks to either ambient (370 ppm) or elevated levels of CO 2 (800 ppm) and then planted in the field; seedling growth was followed for the next six years. Exponential nutrient loading increased seedling height, stem diameter and leader growth, with the largest increases in height and leader length occurring in the first three years after outplanting. Carbon loading increased seedling height and leader length, but only in seedlings that had been exponentially nutrient loaded. A combination of carbon and nutrient loading increased shoot height 26%, stem diameter 37% and leader length 40% over trees that received neither treatment. These results demonstrate that the growth enhancement seen under exponential nutrient loading is maintained under field conditions for at least six years. Carbon loading just before outplanting was a useful supplement to nutrient loading, but was ineffective in the absence of nutrient loading

CO Emission in Low Luminosity, HI Rich Galaxies

We present 12CO 1-0 observations of eleven low luminosity M_B > -18), HI--rich dwarf galaxies. Only the three most metal-rich galaxies, with 12+log(O/H) ~ 8.2, are detected. Very deep CO spectra of six extremely metal-poor systems (12+log(O/H) < 7.5) yield only low upper limits on the CO surface brightness, I_CO < 0.1 K km/s. Three of these six have never before been observed in a CO line, while the others now have much more stringent upper limits. For the very low metallicity galaxy Leo A, we do not confirm a previously reported detection in CO, and the limits are consistent with another recent nondetection. We combine these new observations with data from the literature to form a sample of dwarf galaxies which all have CO observations and measured oxygen abundances. No known galaxies with 12+log(O/H) < 7.9 (Z < 0.1 solar) have been detected in CO. Most of the star-forming galaxies with higher (12+log(O/H) > 8.1) metallicities are detected at similar or higher I_CO surface brightnesses. The data are consistent with a strong dependence of the I_CO/M_H_2 = X_CO conversion factor on ambient metallicity. The strikingly low upper limits on some metal-poor galaxies lead us to predict that the conversion factor is non-linear, increasing sharply below approximately 1/10 of the solar metallicity (12+log(O/H) < 7.9).Comment: 25 pages, 4 figures, 3 tables. Accepted for publication in AJ Tables replaced -- now formated for landscape orientatio

Lineage-specific gene radiations underlie the evolution of novel betalain pigmentation in Caryophyllales.

Betalain pigments are unique to the Caryophyllales and structurally and biosynthetically distinct from anthocyanins. Two key enzymes within the betalain synthesis pathway have been identified: 4,5-dioxygenase (DODA) that catalyzes the formation of betalamic acid and CYP76AD1, a cytochrome P450 gene that catalyzes the formation of cyclo-DOPA. We performed phylogenetic analyses to reveal the evolutionary history of the DODA and CYP76AD1 lineages and in the context of an ancestral reconstruction of pigment states we explored the evolution of these genes in relation to the complex evolution of pigments in Caryophylalles. Duplications within the CYP76AD1 and DODA lineages arose just before the origin of betalain pigmentation in the core Caryophyllales. The duplications gave rise to DODA-α and CYP76AD1-α isoforms that appear specific to betalain synthesis. Both betalain-specific isoforms were then lost or downregulated in the anthocyanic Molluginaceae and Caryophyllaceae. Our findings suggest a single origin of the betalain synthesis pathway, with neofunctionalization following gene duplications in the CYP76AD1 and DODA lineages. Loss of DODA-α and CYP76AD1-α in anthocyanic taxa suggests that betalain pigmentation has been lost twice in Caryophyllales, and exclusion of betalain pigments from anthocyanic taxa is mediated through gene loss or downregulation. [Correction added after online publication 13 May 2015: in the last two paragraphs of the Summary the gene name CYP761A was changed to CYP76AD1.].S.C. was supported by a grant to IRRI from the Bill and Melinda Gates Foundation and UKAID. This work was supported by a National Science Foundation award (grant numbers DEB 1354048 and DEB 1352907) to S.F.B., M.J.M. and S.A.S., and a NERC Independent Research Fellowship to S.F.B. The 1000 Plants (1KP) initiative, led by G.K.S.W., is funded by the Alberta Ministry of Enterprise and Advanced Education, Alberta Innovates Technology Futures (AITF), Innovates Centre of Research Excellence (iCORE), Musea Ventures and BGI-Shenzhen.This is the final version of the article. It first appeared from Wiley via http://dx.doi.org/10.1111/nph.1344

RNA-Seq based phylogeny recapitulates previous phylogeny of the genus Flaveria (Asteraceae) with some modifications

Abstract Background The genus Flaveria has been extensively used as a model to study the evolution of C4 photosynthesis as it contains C3 and C4 species as well as a number of species that exhibit intermediate types of photosynthesis. The current phylogenetic tree of the genus Flaveria contains 21 of the 23 known Flaveria species and has been previously constructed using a combination of morphological data and three non-coding DNA sequences (nuclear encoded ETS, ITS and chloroplast encoded trnL-F). Results Here we developed a new strategy to update the phylogenetic tree of 16 Flaveria species based on RNA-Seq data. The updated phylogeny is largely congruent with the previously published tree but with some modifications. We propose that the data collection method provided in this study can be used as a generic method for phylogenetic tree reconstruction if the target species has no genomic information. We also showed that a “F. pringlei” genotype recently used in a number of labs may be a hybrid between F. pringlei (C3) and F. angustifolia (C3-C4). Conclusions We propose that the new strategy of obtaining phylogenetic sequences outlined in this study can be used to construct robust trees in a larger number of taxa. The updated Flaveria phylogenetic tree also supports a hypothesis of stepwise and parallel evolution of C4 photosynthesis in the Flavaria clade

Multiple photosynthetic transitions, polyploidy, and lateral gene transfer in the grass subtribe Neurachninae

The Neurachninae is the only grass lineage known to contain C3, C4, and C3–C4 intermediate species, and as such has been suggested as a model system for studies of photosynthetic pathway evolution in the Poaceae; however, a lack of a robust phylogenetic framework has hindered this possibility. In this study, plastid and nuclear markers were used to reconstruct evolutionary relationships among Neurachninae species. In addition, photosynthetic types were determined with carbon isotope ratios, and genome sizes with flow cytometry. A high frequency of autopolyploidy was found in the Neurachninae, including in Neurachne munroi F.Muell. and Paraneurachne muelleri S.T.Blake, which independently evolved C4 photosynthesis. Phylogenetic analyses also showed that following their separate C4 origins, these two taxa exchanged a gene encoding the C4 form of phosphoenolpyruvate carboxylase. The C3–C4 intermediate Neurachne minor S.T.Blake is phylogenetically distinct from the two C4 lineages, indicating that intermediacy in this species evolved separately from transitional stages preceding C4 origins. The Neurachninae shows a substantial capacity to evolve new photosynthetic pathways repeatedly. Enablers of these transitions might include anatomical pre-conditions in the C3 ancestor, and frequent autopolyploidization. Transfer of key C4 genetic elements between independently evolved C4 taxa may have also facilitated a rapid adaptation of photosynthesis in these grasses that had to survive in the harsh climate appearing during the late Pliocene in Australia

RNA-Seq based phylogeny recapitulates previous phylogeny of the genus Flaveria (Asteraceae) with some modifications.

BACKGROUND: The genus Flaveria has been extensively used as a model to study the evolution of C4 photosynthesis as it contains C3 and C4 species as well as a number of species that exhibit intermediate types of photosynthesis. The current phylogenetic tree of the genus Flaveria contains 21 of the 23 known Flaveria species and has been previously constructed using a combination of morphological data and three non-coding DNA sequences (nuclear encoded ETS, ITS and chloroplast encoded trnL-F). RESULTS: Here we developed a new strategy to update the phylogenetic tree of 16 Flaveria species based on RNA-Seq data. The updated phylogeny is largely congruent with the previously published tree but with some modifications. We propose that the data collection method provided in this study can be used as a generic method for phylogenetic tree reconstruction if the target species has no genomic information. We also showed that a "F. pringlei" genotype recently used in a number of labs may be a hybrid between F. pringlei (C3) and F. angustifolia (C3-C4). CONCLUSIONS: We propose that the new strategy of obtaining phylogenetic sequences outlined in this study can be used to construct robust trees in a larger number of taxa. The updated Flaveria phylogenetic tree also supports a hypothesis of stepwise and parallel evolution of C4 photosynthesis in the Flavaria clade

A Direct Detection of Dust in the Outer Disks of Nearby Galaxies

We measure the extent of 100 micron galactic emission in two independent galaxy samples using the IRAS 100 micron Sky Survey images and constrain the distribution of dust at large (\ltsim30 kpc) radii. The first sample consists of 90 nearby (v < 6000 km/s) galaxies from the RC3 catalog with similar angular sizes and absolute luminosities (5 arcmin $\leq$D$_{25} \leq$ 10 arcmin and $-22.5 \leq$M$_{B} \leq -18$) that are isolated in the 100 micron images. The second sample consists of 24 local galaxies (v < 1500 km/s, 10 arcmin $\leq$D$_{25} \leq$ 30 arcmin). We rescale the 100 micron images of these galaxies using their optical diameters, D$_{25}$, rotate the images using their optical major axis position angle, construct the mean and median image, and rebin the final images into polar coordinates to study the 100 micron emission as a function of radius and azimuthal angle. We find that the 100 micron emission extends at least to radii of 27 kpc (2$\sigma$ detection) for the typical galaxy in the 5 arcmin - 10 arcmin sample and to 21 kpc (2$\sigma$ detection) in the 10 arcmin - 30 arcmin sample (H$_{0} =$75 km/s/Mpc). In both samples, the emission is preferentially elongated along the optical major axis. We fit an exponential to the 100 micron emission along the major axis and measure a scale length of $2.5 \pm$ 0.8 kpc (90% confidence interval). Using a simple model that relates the far-IR emission to the stellar distribution, we examine the range of acceptable dust mass distributions allowed by our data and conclude that the dust is more extended than the starlight.Comment: 31 pages, 9 figures, AASTEX (aaspp4), accepted for publication in the Astronomical Journa