Phytochrome A elevates plant circadian-clock components to suppress shade avoidance in deep-canopy shade

Shade-avoiding plants can detect the presence of neighboring vegetation and evoke escape responses before canopy cover limits photosynthesis. Rapid stem elongation facilitates light foraging and enables plants to overtop competitors. A major regulator of this response is the phytochrome B photoreceptor, which becomes inactivated in light environments with a low ratio of red to far-red light (low R:FR), characteristic of vegetational shade. Although shade avoidance can provide plants with a competitive advantage in fast-growing stands, excessive stem elongation can be detrimental to plant survival. As such, plants have evolved multiple feedback mechanisms to attenuate shade-avoidance signaling. The very low R:FR and reduced levels of photosynthetically active radiation (PAR) present in deep canopy shade can, together, trigger phytochrome A (phyA) signaling, inhibiting shade avoidance and promoting plant survival when resources are severely limited. The molecular mechanisms underlying this response have not been fully elucidated. Here, we show that Arabidopsis thaliana phyA elevates early-evening expression of the central circadian-clock components TIMING OF CAB EXPRESSION 1 (TOC1), PSEUDO RESPONSE REGULATOR 7 (PRR7), EARLY FLOWERING 3 (ELF3), and ELF4 in photocycles of low R:FR and low PAR. These collectively suppress stem elongation, antagonizing shade avoidance in deep canopy shade.

Shade Avoidance and Light Foraging of a Clonal Woody Species, Pachysandra terminalis

(1) Background: A central subject in clonal plant ecology is to elucidate the mechanism by which clones forage resources in heterogeneous environments. Compared with studies conducted in laboratories or experimental gardens, studies on light foraging of forest woody clonal plants in their natural habitats are limited. (2) Methods: We investigated wild populations of an evergreen clonal understory shrub, Japanese pachysandra (Pachysandra terminalis Siebold & Zucc.), in two cool-temperate forests in Japan. (3) Results: Similar to the results of herbaceous clonal species, this species formed a dense stand in a relatively well-lit place, and a sparse stand in a shaded place. Higher specific rhizome length (i.e., length per unit mass) in shade resulted in lower ramet population density in shade. The individual leaf area, whole-ramet leaf area, or ramet height did not increase with increased light availability. The number of flower buds per flowering ramet increased as the canopy openness or population density increased. (4) Conclusions: Our results provide the first empirical evidence of shade avoidance and light foraging with morphological plasticity for a clonal woody species.

Morphometric comparisons and novel observations of diurnal and low-light-foraging bees

Low-light adapted bees are substantially understudied components of the bee fauna, particularly in Australia. Whilst several species in Australia are thought to be adapted to low-light conditions, explicit records of these taxa actually foraging at twilight or night are absent from the scientific literature. We present the first observations of Australian bees foraging in low-light conditions as well as the first evidence of low-light foraging behaviour in the colletid bee subfamily, Hylaeinae. Using morphometrics of Australian and more broadly-distributed diurnal, facultative low-light and obligate low-light adapted bees, we explore the use of morphological traits to objectively assess possible low-light behaviour and corroborate low-light collection events. Our results show that it is possible to morphologically distinguish between diurnal and low-light adapted bees, and that there is a spectrum of characters that are associated with low light conditions. We use GIS to show that low-light adapted species occur mostly in the tropics, but that some species have subtropical, arid and even temperate distributions. As low-light foraging behaviour in bees is infrequently reported, it appears that low-light foraging behaviour is more common than currently appreciated, highlighting the need for extended bee-sampling periods and more consistent collection data to increase the understanding of this little-understood aspect of bee behaviour.

Species and sex differences in eye morphometry and visual responsivity of two crepuscular sweat bee species (Megalopta spp., Hymenoptera: Halictidae)

Visually dependent dim-light foraging has evolved repeatedly, broadening the ecological niches of some species. Many dim-light foraging lineages evolved from diurnal ancestors, requiring immense visual sensitivity increases to compensate for light levels a billion times dimmer than daylight. Some taxa, such as bees, are anatomically constrained by apposition compound eyes, which function well in daylight but not in starlight. Even with this constraint, the bee genus Megalopta has incredibly sensitive eyes, foraging in light levels up to nine orders of magnitude dimmer than diurnal relatives. Despite many behavioural studies, variation in visual sensitivity and eye morphometry has not been investigated within and across Megalopta species. Here we quantify external eye morphology (corneal area and facet size) for sympatric species of Megalopta, M. genalis and M. amoena, which forage during twilight. We use electroretinograms to show that males, despite being smaller than females, have equivalent visual sensitivity and increased retinal responsivity. Although males have relatively larger eyes compared with females, corneal area and facet size were not correlated with retinal responsivity, suggesting that males have additional non-morphological adaptations to increase retinal responsiveness. These findings provide the foundation for future work into the neural and physiological mechanisms that interface with morphology to influence visual sensitivity, with implications for understanding niche exploitation.

Photic niche invasions: phylogenetic history of the dim-light foraging augochlorine bees (Halictidae)

Most bees rely on flowering plants and hence are diurnal foragers. From this ancestral state, dim-light foraging in bees requires significant adaptations to a new photic environment. We used DNA sequences to evaluate the phylogenetic history of the most diverse clade of Apoidea that is adapted to dim-light environments (Augochlorini: Megalopta , Megaloptidia and Megommation ). The most speciose lineage, Megalopta , is distal to the remaining dim-light genera, and its closest diurnal relative ( Xenochlora ) is recovered as a lineage that has secondarily reverted to diurnal foraging. Tests for adaptive protein evolution indicate that long-wavelength opsin shows strong evidence of stabilizing selection, with no more than five codons (2%) under positive selection, depending on analytical procedure. In the branch leading to Megalopta , the amino acid of the single positively selected codon is conserved among ancestral Halictidae examined, and is homologous to codons known to influence molecular structure at the chromophore-binding pocket. Theoretically, such mutations can shift photopigment λ max sensitivity and enable visual transduction in alternate photic environments. Results are discussed in light of the available evidence on photopigment structure, morphological specialization and biogeographic distributions over geological time.