C4 photosynthesis is a complex trait that boosts productivity in tropical conditions. Compared with C3 species, the C4 state seems to require numerous novelties, but species comparisons can be confounded by long divergence times. Here, we exploit the photosynthetic diversity that exists within a single species, the grass Alloteropsis semialata, to detect changes in gene expression associated with different photosynthetic phenotypes. Phylogenetically informed comparative transcriptomics show that intermediates with a weak C4 cycle are separated from the C3 phenotype by increases in the expression of 58 genes (0.22% of genes expressed in the leaves), including those encoding just three core C4 enzymes: aspartate aminotransferase, phosphoenolpyruvate carboxykinase, and phosphoenolpyruvate carboxylase. The subsequent transition to full C4 physiology was accompanied by increases in another 15 genes (0.06%), including only the core C4 enzyme pyruvate orthophosphate dikinase. These changes probably created a rudimentary C4 physiology, and isolated populations subsequently improved this emerging C4 physiology, resulting in a patchwork of expression for some C4 accessory genes. Our work shows how C4 assembly in A. semialata happened in incremental steps, each requiring few alterations over the previous step. These create short bridges across adaptive landscapes that probably facilitated the recurrent origins of C4 photosynthesis through a gradual process of evolution.
C4 photosynthesis is a complex trait that boosts productivity in warm environments. Paradoxically, it evolved independently in numerous plant lineages, despite requiring specialised leaf anatomy. The anatomical modifications underlying C4 evolution have previously been evaluated through interspecific comparisons, which capture numerous changes besides those needed for C4 functionality. Here, we quantify the anatomical changes accompanying the transition between non‐C4 and C4 phenotypes by sampling widely across the continuum of leaf anatomical traits in the grass Alloteropsis semialata. Within this species, the only trait that is shared among and specific to C4 individuals is an increase in vein density, driven specifically by minor vein development that yields multiple secondary effects facilitating C4 function. For species with the necessary anatomical preconditions, developmental proliferation of veins can therefore be sufficient to produce a functional C4 leaf anatomy, creating an evolutionary entry point to complex C4 syndromes that can become more specialised.
Three species of Nassella have naturalized in South Africa. Nassella trichotoma and N. tenuissima are declared weeds under category 1b of the National Environmental Management: Biodiversity Act (NEM:BA) and occur mainly in the montane grasslands of the Western and Eastern Cape provinces. Nassella neesiana is not listed in NEM:BA but is naturalized in the Eastern Cape, Western Cape and Free State provinces. Research conducted in the 1970s and 1980s led to vigorous government-funded awareness and control campaigns which ended in 2000. No research on Nassella distribution or control has been undertaken since then. Despite this hiatus, Nassella remains a dangerous genus in southern Africa, given the serious impacts of these species in similar social-ecological systems in Australia and New Zealand. This paper presents a synthesis of available information about Nassella invasions in South Africa and identifies research gaps. It specifically addresses these questions: What identification issues exist? What is the current spatial distribution of Nassella? What is the autecology of the genus? What are the social-ecological impacts of Nassella? What control measures are currently applied and what are their strengths and limitations? What do we know about Nassella distribution and its response to climate change? This paper highlights many knowledge gaps about Nassella, such as the species' current distribution range, field identification and detection difficulties, and the uncoordinated control efforts that require urgent research to inform an effective management response.
The first comprehensive plant checklist for the Bvumba massif, situated in the Manica Highlands along the Zimbabwe-Mozambique border, is presented. Although covering only 276 km2, the flora is rich with 1250 taxa (1127 native taxa and 123 naturalised introductions). There is a high proportion of Orchidaceae and Pteridophyta, with both groups showing a higher richness than for adjacent montane areas, which may be due to the massif’s relatively high moisture levels as a result of frequent cloud cover. However, in contrast to other mesic montane regions in southern Africa, there are relatively few near-endemic or range-restricted taxa: there is only one local endemic, Aeranthes africana, an epiphytic forest orchid. This is likely to be an effect of the massif having limited natural grassland compared to forest, the former being the most endemic-rich habitat in southern African mountains outside of the Fynbos Biome. Six other near-endemic taxa with limited distribution in this portion of the Manica Highlands are highlighted. The high number of invasive species is probably a result of diverse human activities in the area. The main species of concern are Acacia melanoxylon, a tree that is invading grassland and previously cultivated land, the forest herb Hedychium gardnerianum which in places is transforming forest understorey with an adverse effect on some forest birds, and the woody herb Vernonanthura polyanthes which invades cleared forest areas after fire. Future botanical work in the massif should focus on a more detailed exploration of the poorly known Serra Vumba on the Mozambican side and on the drier western slopes. This will allow for a more detailed analysis of patterns of endemism across the Manica Highlands.
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