Howard Hughes Medical Institute, University of California, Berkeley, CA 94720.
Department of Plant and Microbial Biology, University of California, Berkeley, CA 94720.
Proc Natl Acad Sci U S A. 2017 Aug 15;114(33):E7002-E7008. doi: 10.1073/pnas.1704373114. Epub 2017 Jul 31.
Although sunlight provides the energy necessary for plants to survive and grow, light can also damage reaction centers of photosystem II (PSII) and reduce photochemical efficiency. To prevent damage, plants possess photoprotective mechanisms that dissipate excess excitation. A subset of these mechanisms is collectively referred to as NPQ, or nonphotochemical quenching of chlorophyll fluorescence. The regulation of NPQ is intrinsically linked to the cycling of xanthophylls that affects the kinetics and extent of the photoprotective response. The violaxanthin cycle (VAZ cycle) and the lutein epoxide cycle (LxL cycle) are two xanthophyll cycles found in vascular plants. The VAZ cycle has been studied extensively, owing in large part to its presence in model plant species where mutants are available to aid in its characterization. In contrast, the LxL cycle is not found in model plants, and its role in photosynthetic processes has been more difficult to define. To address this challenge, we introduced the LxL cycle into and functionally isolated it from the VAZ cycle. Using these plant lines, we showed an increase in dark-acclimated PSII efficiency associated with Lx accumulation and demonstrated that violaxanthin deepoxidase is responsible for the light-driven deepoxidation of Lx. Conversion of Lx to L was reversible during periods of low light and occurred considerably faster than rates previously described in nonmodel species. Finally, we present clear evidence of the LxL cycle's role in modulating a rapid component of NPQ that is necessary to prevent photoinhibition in excess light.
虽然阳光为植物的生存和生长提供了所需的能量,但光也会破坏光系统 II (PSII) 的反应中心,降低光化学效率。为了防止损伤,植物具有防止光破坏的机制,可以耗散过量的激发能。这些机制中的一部分被统称为 NPQ,即叶绿素荧光的非光化学猝灭。NPQ 的调节与叶黄素的循环密切相关,叶黄素的循环会影响光保护反应的动力学和程度。 violaxanthin 循环 (VAZ 循环) 和 lutein epoxide cycle (LxL cycle) 是在维管植物中发现的两种叶黄素循环。VAZ 循环已经被广泛研究,这在很大程度上是由于其存在于模式植物物种中,这些物种有突变体可供研究以帮助其特性的描述。相比之下,LxL 循环在模式植物中不存在,其在光合作用过程中的作用更难定义。为了解决这个挑战,我们将 LxL 循环引入到 中,并将其与 VAZ 循环功能上分离。利用这些植物品系,我们发现 Lx 的积累与暗适应 PSII 效率的增加有关,并证明了 violaxanthin deepoxidase 负责 Lx 的光驱动深氧化。在低光条件下,Lx 转化为 L 是可逆的,其速度比以前在非模式物种中描述的速度快得多。最后,我们提供了明确的证据表明,LxL 循环在调节快速的 NPQ 组分中起作用,这对于防止过量光下的光抑制是必要的。