Summary
Nitrogen (N) deficiency triggers an accumulation of a storage lipid triacylglycerol (TAG) in seed plants and algae. Whereas the metabolic pathway and regulatory mechanism to synthesize TAG from diacylglycerol are well known, enzymes involved in the supply of diacylglycerol remain elusive under N starvation.
Lysophosphatidic acid acyltransferase (LPAT) catalyzes an important step of the de novo phospholipid biosynthesis pathway and thus has a strong flux control in the biosynthesis of phospholipids and TAG. Five LPAT isoforms are known in Arabidopsis; however, the functions of LPAT4 and LPAT5 remain elusive.
Here, we show that LPAT4 and LPAT5 are functional endoplasmic‐reticulum‐localized LPATs. Seedlings of the double knockout mutant lpat4‐1 lpat5‐1 showed reduced content of phospholipids and TAG under normal growth condition. Under N starvation, lpat4‐1 lpat5‐1 seedlings showed severer growth defect than the wild‐type in shoot. The phenotype was similar to dgat1‐4, which affects a major TAG biosynthesis pathway and showed similarly reduced TAG content as the lpat4‐1 lpat5‐1.
We suggest that LPAT4 and LPAT5 may redundantly function in endoplasmic‐reticulum‐localized de novo glycerolipid biosynthesis for phospholipids and TAG, which is important for the N starvation response in Arabidopsis.
Periods of drought, that threaten crop production, are expected to become more prominent in large parts of the world, making it necessary to explore all aspects of plant growth and development, to breed, modify and select crops adapted to such conditions. One such aspect is the xylem, where influencing the size and number of the water-transporting xylem vessels, may impact on hydraulic conductance and drought tolerance. Here, we focus on how plants adjust their root xylem as a response to reduced water availability. While xylem response has been observed in a wide array of species, most of our knowledge on the molecular mechanisms underlying xylem plasticity comes from studies on the model plant Arabidopsis thaliana. When grown under water limiting conditions, Arabidopsis rapidly adjusts its development to produce more xylem strands with altered identity in an abscisic acid (ABA) dependent manner. Other hormones such as auxin and cytokinin are essential for vascular patterning and differentiation. Their balance can be perturbed by stress, as evidenced by the effects of enhanced jasmonic acid signaling, which results in similar xylem developmental alterations as enhanced ABA signaling. Furthermore, brassinosteroids and other signaling molecules involved in drought tolerance can also impact xylem development. Hence, a multitude of signals affect root xylem properties and, potentially, influence survival under water limiting conditions. Here, we review the likely entangled signals that govern root vascular development, and discuss the importance of taking root anatomical traits into account when breeding crops for enhanced resilience toward changes in water availability.
Summary
Unsaturation of membrane glycerolipid classes at their hydrophobic fatty acid tails critically affects the physical nature of the lipid molecule. In Arabidopsis thaliana, 7 fatty acid desaturases (FADs) differently desaturate each glycerolipid class in plastids and the endoplasmic reticulum (ER). Here, we showed that polyunsaturation of ER glycerolipids is required for the ER stress response. Through systematic screening of FAD mutants, we found that a mutant of FAD2 resulted in a hypersensitive response to tunicamycin, a chemical inducer of ER stress. FAD2 converts oleic acid to linoleic acid of the fatty acyl groups of ER‐synthesized phospholipids. Our functional in vivo reporter assay revealed the ER localization and distinct tissue‐specific expression patterns of FAD2. Moreover, glycerolipid profiling of both mutants and overexpressors of FAD2 under tunicamycin‐induced ER stress conditions, along with phenotypic screening of the mutants of the FAD family, suggested that the ratio of monounsaturated fatty acids to polyunsaturated fatty acids, particularly 18:1 to 18:2 species, may be an important factor in allowing the ER membrane to cope with ER stress. Therefore, our results suggest that membrane lipid polyunsaturation mediated by FAD2 is involved in ER stress tolerance in Arabidopsis.
Upon phosphate starvation, plants retard shoot growth but promote root development presumably to enhance phosphate assimilation from the ground. Membrane lipid remodelling is a metabolic adaptation that replaces membrane phospholipids by non-phosphorous galactolipids, thereby allowing plants to obtain scarce phosphate yet maintain the membrane structure. However, stoichiometry of this phospholipid-to-galactolipid conversion may not account for the massive demand of membrane lipids that enables active growth of roots under phosphate starvation, thereby suggesting the involvement of de novo phospholipid biosynthesis, which is not represented in the current model. We overexpressed an endoplasmic reticulum-localized lysophosphatidic acid acyltransferase, LPAT2, a key enzyme that catalyses the last step of de novo phospholipid biosynthesis. Two independent LPAT2 overexpression lines showed no visible phenotype under normal conditions but showed increased root length under phosphate starvation, with no effect on phosphate starvation response including marker gene expression, root hair development and anthocyanin accumulation. Accompanying membrane glycerolipid profiling of LPAT2-overexpressing plants revealed an increased content of major phospholipid classes and distinct responses to phosphate starvation between shoot and root. The findings propose a revised model of membrane lipid remodelling, in which de novo phospholipid biosynthesis mediated by LPAT2 contributes significantly to root development under phosphate starvation.
Phosphorus is essential for plant viability. Phosphate-starved plants trigger membrane lipid remodeling to replace membrane phospholipids by non-phosphorus galactolipids presumably to acquire scarce phosphate source. Phosphoethanolamine/phosphocholine phosphatase 1 (PECP1) and phosphate starvation-induced gene 2 (PS2) belong to an emerging class of phosphatase induced by phosphate starvation and dephosphorylates phosphocholine and phosphoethanolamine (PEtn)
in vivo
. However, detailed spatiotemporal expression pattern as well as subcellular localization has not been investigated yet. Here, by constructing transgenic plants harboring functional translational promoter–reporter fusion system, we showed the expression pattern of PECP1 and PS2 in different tissues and in response to phosphate starvation. Besides, the Venus fluorescent reporter revealed that both are localized at the ER. Characterization of transgenic plants that overexpress PECP1 or PS2 showed that their activity toward PEtn may be different
in vivo
. We suggest that PECP1 and PS2 are ER-localized phosphatases that show similar expression pattern yet have a distinct substrate specificity
in vivo.
Non-specific phospholipase Cs (NPCs) are responsible for membrane lipid remodeling that involves hydrolysis of the polar head group of membrane phospholipids. Arabidopsis NPC2 and NPC6 are essential in gametogenesis, but their underlying role in the lipid remodeling remains elusive. Here, we show that these NPCs are required for triacylglycerol (TAG) production in pollen tube growth. NPC2 and NPC6 are highly expressed in developing pollen tubes and are localized at the endoplasmic reticulum. Mutants of NPC2 and NPC6 showed reduced rate of pollen germination, length of pollen tube and amount of lipid droplets (LDs). Overexpression of NPC2 or NPC6 induced LD accumulation, which suggests that these NPCs are involved in LD production. Furthermore, mutants defective in the biosynthesis of TAG, a major component of LDs, showed defective pollen tube growth. These results suggest that NPC2 and NPC6 are essential in gametogenesis for a role in hydrolyzing phospholipids and producing TAG required for pollen tube growth. Thus, lipid remodeling from phospholipids to TAG during pollen tube growth represents an emerging role for the NPC family in plant developmental control.
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