
Surviving in the understorey: ๐๐บ๐ค๐ญ๐ข๐ฎ๐ฆ๐ฏ ๐ฑ๐ถ๐ณ๐ฑ๐ถ๐ณ๐ข๐ด๐ค๐ฆ๐ฏ๐ด in dynamic light
Variegated leaves are common in forest understorey species. As sunflecks (patches of direct sunlight) pass over the leaves the plant is exposed to rapid, dynamic changes between deep shade and high intensity light. The different zones of the leaf interact differently with high or low light intensities. The pale regions perform photosynthesis best at high light (in sunflecks) and the dark regions maximise photosynthesis in the shade. In this way some part of the leaf can perform photosynthesis well no matter how the light changes.
In this example, purple cyclamen (๐๐บ๐ค๐ญ๐ข๐ฎ๐ฆ๐ฏ ๐ฑ๐ถ๐ณ๐ฑ๐ถ๐ณ๐ข๐ด๐ค๐ฆ๐ฏ๐ด: Primulaceae) grows in the understorey of woodland at Monte Barro, Italy. ๐๐บ๐ค๐ญ๐ข๐ฎ๐ฆ๐ฏ ๐ฑ๐ถ๐ณ๐ฑ๐ถ๐ณ๐ข๐ด๐ค๐ฆ๐ฏ๐ด has two regions of the leaf that interact differently with light. The pale regions have air spaces under the outer, epidermal, layer which form a reflective โblisterโ (inside the leaf, the cells are rounder, letting air pass around them). This is associated with low chlorophyll pigment contents and also a lack of red pigment (anthocyanin) in the lower epidermis. This pale region performs photosynthesis most effectively when direct, bright, white sunlight passes over the leaf, reflecting excess light or letting it pass through, and leaving just enough for effective photosynthesis. However, in the shade the pale regions do not have enough chlorophyll to scavenge light, and photosynthesis is minimal. The dark areas have closely packed cells with high chlorophyll contents, which can intercept the scarce photons of light that are available in the shade. Also, the cells of the lower epidermis in these regions have high anthocyanin contents, which reflects red light back up into the leaf, giving the chlorophyll a second chance to absorb it (the โred mirrorโ effect). While these are effective mechanisms in the shade, the dark regions become over-exposed in the sunflecks, with excessive energy being absorbed and hindering photosynthesis (an effect known as photoinhibition). Having two regions of the leaf optimised for high or low light means that there is always some part of the leaf that is able to photosynthesise, even though the light changes throughout the day or under different weather conditions.
๐ง๐ต๐ถ๐ ๐ฐ๐ต๐ฎ๐ป๐ป๐ฒ๐น ๐ถ๐ ๐ป๐ผ๐ ๐บ๐ผ๐ป๐ฒ๐๐ถ๐๐ฒ๐ฑ.
๐ฆ๐ฐ๐ถ๐ฒ๐ป๐๐ถ๐ณ๐ถ๐ฐ ๐น๐ถ๐๐ฒ๐ฟ๐ฎ๐๐๐ฟ๐ฒ:
Klanฤnik, K., Levpuลกฤek, M. & Gaberลกฤik, A. (2016) Variegation and red abaxial epidermis define the leaf optical properties of ๐๐บ๐ค๐ญ๐ข๐ฎ๐ฆ๐ฏ ๐ฑ๐ถ๐ณ๐ฑ๐ถ๐ณ๐ข๐ด๐ค๐ฆ๐ฏ๐ด. Flora, 224: 87-95. https://doi.org/10.1016/j.flora.2016....
Pierce, S. (2003) Variegated foliage and photosynthesis. Journal of the Bromeliad Society, 53(5): 216-240. https://www.researchgate.net/publicat...
Zhang, J.-H., Zeng, J.-C., Wang, X.-M., Chen, S.-F., Albach, D.C., Li, H.-Q. (2020) A revised classification of leaf variegation types. Flora, 272: 151703. https://doi.org/10.1016/j.flora.2020..... https://tiantong.ecnu.edu.cn/kindedit...
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