甘油-3-磷酸酰基转移酶（Glycerol-3-phosphate acyltransferase，GPAT）作为TAG合成通路中的第一个酶和重要限速酶，在植物生长发育以及不同生长阶段的环境适应中发挥重要作用。拟南芥内质网定位的GPATs包括GPAT4-9，其中AtGPAT9被报道参与三酰甘油（TAG）的生物合成，影响种子中油的含量。GPAT4-8仅被报道出参与角质以及软木脂等植物脂质的合成。然而，内质网定位的GPAT4-8蛋白是否参与脂滴（Lipid Droplet）生成，且其调控的机制尚不清楚。本论文主要以拟南芥GPAT4/8为主要研究对象，研究了内质网定位的GPAT4/8复合体对热胁迫、盐胁迫以及ABA响应的调控作用，同时初步确定了GPAT4/8与ATI1/2（ATG8相互作用蛋白1/2）相互作用，共同调控热胁迫下的LD增殖的分子机理。主要结果如下：

拟南芥突变体在200 mM NaCl处理后，gpat4 gpat8双突变体子叶白化死亡率显著高于WT；在45℃高温处理后，仅有gpat4 gpat8双突变体子叶以及四周龄植株死亡率显著高于WT，其他双突变体无显著差异；在脱落酸（ABA）处理下，gpat4 gpat8双突变体的相对主根伸长量表现出敏感表型。上述结果表明，GPAT4和GPAT8参与调控植物盐胁迫、高温响应以及ABA信号通路。

利用生信分析网站AlphaFold3预测到GPAT4/6/8与ATI1/2之间可能存在相互作用。在酵母双杂交体系中，明确了内质网GPAT4-9中仅有GPAT4/6/8与ATI1/2存在相互作用。利用荧光素酶互补实验（Luciferase complementation imaging，LCI）、Pull-down、以及能量共振转移荧光寿命成像（Förster resonance energy transfer by fluorescence lifetime imaging，FRET-FLIM）等技术手段进一步验证了它们体内与体外存在互作，同时GPAT4/6/8发夹结构HP区域是其主要的互作区域，而ATI1/2主要互作区域是N端序列。

研究发现，黑暗周期结束后，gpat4 gpat8与ati1 ati2双突变体材料叶片LD丰度较WT显著降低。热胁迫处理后，与WT相比，gpat4 gpat8以及ati1 ati2双突变体的死亡率表型显著增加，gpat4 gpat8双突变体叶片的LD丰度较WT显著降低，表明GPAT4/8与ATI1/2参与高温胁迫下的脂滴合成过程。利用GPAT4/6/8-mCherry转基因植株，观察发现荧光在光照条件仅定位在内质网中，而油酸或热处理后，其定位由内质网靶向脂滴表面，暗示其参与脂滴的增殖过程；GFP-ATI2表现类似的由内质网靶向合成的脂滴表面特征。利用溶血磷脂酸（Lysophosphatidic acid，LPA）的荧光探针GFP-GLP₁₅₁观察发现，LPA也可以从内质网靶向脂滴表面，且荧光漂白恢复实验后其荧光强度又立即恢复，表明LPA可能参与脂滴的合成过程。

综上所述，本研究初步揭示了GPAT4/8及其产物LPA参与调控脂滴合成与高温响应的分子机理。在高温胁迫下，GPAT4/8与ATI1/2形成复合体，共同由内质网移动到脂滴表面，调控LPA的合成，通过某种机制调控脂滴的合成或代谢，为后期深入研究脂滴增殖及植物应答高温胁迫提供了重要理论依据。

Glycerol-3-phosphate acyltransferase (GPAT), as the first enzyme and an important rate-limiting enzyme in the TAG synthesis pathway, plays an important role in plant growth and development and environmental adaptation at different growth stages. GPATs located in the endoplasmic reticulum of Arabidopsis thaliana include GPAT4-9.AtGPAT9 has been reported to be involved in the biosynthesis of triacylglycerol (TAG) and affect the oil content in seeds. GPAT4-8 has only been reported to be involved in the synthesis of plant lipids such as cutin and cork. However, whether the endoplasmic reticulum-localized GPAT4-8 protein is involved in LD formation and the mechanism of its regulation is still unclear. In this paper, GPAT4/8 was used as the main research object to study the regulation of GPAT4/8 complex located in endoplasmic reticulum on heat stress, salt stress and ABA response. At the same time, the molecular mechanism of GPAT4 / 8 interacting with ATI1/2 (ATG8 interacting protein 1/2) to regulate LD proliferation under heat stress was preliminarily determined. The main results are as follows :

After 200 mM NaCl treatment, the cotyledon albino mortality of gpat4 gpat8 double mutant was significantly higher than that of WT. After high temperature treatment at 45℃, only the mortality of cotyledons and four-week-old plants of gpat4 gpat8 double mutants was significantly higher than that of WT, and there was no significant difference in other double mutants. Under abscisic acid (ABA) treatment, the relative main root elongation of gpat4 gpat8 double mutant showed a sensitive phenotype. The above results indicate that GPAT4 and GPAT8 are involved in the regulation of plant salt stress, high temperature response and ABA signaling pathway.

The interaction between GPAT4/6/8 and ATI1/2 was predicted by AlphaFold3. In the yeast two-hybrid system, it was clear that only GPAT4/6/8 interacted with ATI1/2 in the endoplasmic reticulum GPAT4-9. Luciferase complementation imaging (LCI), Pull-down, and Förster resonance energy transfer by fluorescence lifetime imaging (FRET-FLIM) were used to further verify their interaction in vivo and in vitro. At the same time, the HP region of GPAT4/6/8 hairpin structure is the main interaction region, while the main interaction region of ATI1/2 is the N-terminal sequence.

It was found that the LD abundance of gpat4 gpat8 and ati1 ati2 double mutant leaves was significantly lower than that of WT after the end of the dark cycle. After heat stress treatment, compared with WT, the mortality phenotype of gpat4 gpat8 and ati1 ati2 double mutants was significantly increased, and the LD abundance of gpat4 gpat8 double mutant leaves was significantly lower than that of WT, indicating that GPAT4/8 and ATI1/2 were involved in the process of lipid droplet proliferation under high temperature stress. Using GPAT4/6/8-mCherry transgenic plants, it was observed that fluorescence was only localized in the endoplasmic reticulum under light conditions, while after oleic acid or heat treatment, its localization was targeted by the endoplasmic reticulum to the surface of lipid droplets, suggesting that it was involved in the proliferation of lipid droplets. GFP-ATI2 showed similar surface characteristics of lipid droplets targeted by endoplasmic reticulum. Using the fluorescent probe GFP-GLP₁₅₁ of LPA, it was found that LPA could also target the surface of lipid droplets from the endoplasmic reticulum, and its fluorescence intensity recovered immediately after the fluorescence bleaching recovery experiment, indicating that LPA may be involved in the proliferation of lipid droplets.

In summary, this study preliminarily revealed the molecular mechanism of GPAT4/8 and its product LPA involved in regulating lipid droplet synthesis and high temperature response. Under high temperature stress, GPAT4/8 and ATI1/2 form a complex, moving together from the endoplasmic reticulum to the surface of lipid droplets, regulating the synthesis of LPA, regulating the synthesis or metabolism of lipid droplets through a certain mechanism, which provides an important theoretical basis for further study of lipid droplet proliferation and plant response to high temperature stress.

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