Amiard Véronique, Demmig-Adams Barbara, Mueh Kristine E, Turgeon Robert, Combs Andrew F, Adams William W
Department of Ecology & Evolutionary Biology, University of Colorado, Boulder, CO 80309-0334, USA.
Present address: Instituto de Investigaciones Agropecuarias, Unidad de Biotecnología, INIA Carillanca, Casilla 58-D, Temuco, Chile.
New Phytol. 2007;173(4):722-731. doi: 10.1111/j.1469-8137.2006.01954.x.
Phloem cells adjacent to sieve elements can possess wall invaginations. The role of light and jasmonic acid signaling in wall ingrowth development was examined in pea companion cells (CCs), Arabidopsis thaliana phloem parenchyma cells (PCs), and in Senecio vulgaris (with ingrowths in both cell types). Features characterized included wall ingrowths (from electron microscopic images), foliar vein density and photosynthetic capacity. In Arabidopsis, wall ingrowths were bulky compared with finger-like invaginations in pea and S. vulgaris. Relative to low light (LL), wall invagination in both CCs and PCs was greater in high light (HL). Treatment with methyl jasmonate in LL had no effect on CCs, but increased PC wall ingrowths. LL-to-HL transfer resulted in significantly less wall ingrowth in the fad7-1 fad8-1 (jasmonate-deficient) Arabidopsis mutant relative to the wild type. These results suggest that chloroplast oxidative status, via chloroplast-derived jasmonates, may modulate phloem structure and function. While CC wall ingrowths facilitate phloem loading by expanding the membrane area available for active uptake, one can speculate that phloem PC ingrowths may have two potential roles: to increase the efflux of sugars and/or protons into the apoplast to augment phloem loading; and/or to protect the phloem against pathogens and/or insects.
与筛管分子相邻的韧皮部细胞可能具有细胞壁内陷。研究了光和茉莉酸信号在豌豆伴胞(CCs)、拟南芥韧皮部薄壁细胞(PCs)以及普通千里光(两种细胞类型均有内陷)细胞壁内突生长发育中的作用。所表征的特征包括细胞壁内突(来自电子显微镜图像)、叶脉密度和光合能力。在拟南芥中,与豌豆和普通千里光中手指状的内陷相比,细胞壁内突较为粗大。相对于弱光(LL),高光(HL)条件下CCs和PCs中的细胞壁内陷都更多。在弱光条件下用茉莉酸甲酯处理对CCs没有影响,但增加了PCs的细胞壁内突。从弱光转移到高光条件下,与野生型相比,茉莉酸缺陷型拟南芥突变体fad7-1 fad8-1中的细胞壁内突显著减少。这些结果表明,叶绿体的氧化状态可能通过叶绿体衍生的茉莉酸来调节韧皮部的结构和功能。虽然CCs的细胞壁内突通过扩大可用于主动吸收的膜面积来促进韧皮部装载,但可以推测韧皮部PCs的内突可能有两个潜在作用:增加糖分和/或质子向质外体的外流以增强韧皮部装载;和/或保护韧皮部免受病原体和/或昆虫侵害。