Instituto Biofisika (CSIC, UPV/EHU), Universidad del País Vasco, Leioa, Spain.
Departamento de Bioquímica, Universidad del País Vasco, Leioa, Spain.
FASEB J. 2021 Jun;35(6):e21657. doi: 10.1096/fj.202001879RR.
Sphingolipids (SL) are ubiquitous in mammalian cell membranes, yet there is little data on the behavior of cells under SL-restriction conditions. LY-B cells derive from a CHO linein whichserine palmitoyl transferase (SPT), thus de novo SL synthesis, is suppressed, while maintaining the capacity of taking up and metabolizing exogenous sphingoid bases from the culture medium. In this study, LY-B cells were adapted to grow in a fetal bovine serum (FBS)-deficient medium to avoid external uptake of lipids. The lowest FBS concentration that allowed LY-B cell growth, though at a slow rate, under our conditions was 0.04%, that is, 250-fold less than the standard (10%) concentration. Cells grown under limiting SL concentrations remained viable for at least 72 hours. Enriching with sphingomyelin the SL-deficient medium allowed the recovery of growth rates analogous to those of control LY-B cells. Studies including whole cells, plasma membrane preparations, and derived lipid vesicles were carried out. Laurdan fluorescence was recorded to measure membrane molecular order, showing a significant decrease in the rigidity of LY-B cells, not only in plasma membrane but also in whole cell lipid extract, as a result of SL limitation in the growth medium. Plasma membrane preparations and whole cell lipid extracts were also studied using atomic force microscopy in the force spectroscopy mode. Force measurements demonstrated that lower breakthrough forces were required to penetrate samples obtained from SL-poor LY-B cells than those obtained from control cells. Mass-spectroscopic analysis was also a helpful tool to understand the rearrangement undergone by the LY-B cell lipid metabolism. The most abundant SL in LY-B cells, sphingomyelin, decreased by about 85% as a result of SL limitation in the medium, the bioactive lipid ceramide and the ganglioside precursor hexosylceramide decreased similarly, together with cholesterol. Quantitative SL analysis showed that a 250-fold reduction in sphingolipid supply to LY-B cells led only to a sixfold decrease in membrane sphingolipids, underlining the resistance to changes in composition of these cells. Plasma membrane compositions exhibited similar changes, at least qualitatively, as the whole cells with SL restriction. A linear correlation was observed between the sphingomyelin concentration in the membranes, the degree of lipid order as measured by laurdan fluorescence, and membrane breakthrough forces assessed by atomic force microscopy. Smaller, though significant, changes were also detected in glycerophospholipids under SL-restriction conditions.
鞘脂类(SL)普遍存在于哺乳动物细胞膜中,但关于细胞在 SL 限制条件下的行为的数据很少。LY-B 细胞来源于 CHO 细胞系,其中丝氨酸棕榈酰转移酶(SPT)被抑制,因此不能从头合成 SL,但仍能从培养基中摄取和代谢外源性鞘氨醇碱基。在这项研究中,LY-B 细胞适应在无胎牛血清(FBS)的培养基中生长,以避免外源性脂质摄取。在我们的条件下,允许 LY-B 细胞生长的最低 FBS 浓度为 0.04%,即比标准(10%)浓度低 250 倍。在限制 SL 浓度下生长的细胞至少能存活 72 小时。在缺乏 SL 的培养基中添加鞘磷脂可以恢复与对照 LY-B 细胞相当的生长速度。进行了包括全细胞、质膜制剂和衍生脂质体在内的研究。使用 Laurdan 荧光测量膜分子的有序性,结果表明,由于生长培养基中 SL 的限制,LY-B 细胞的刚性显著降低,不仅在质膜中,而且在全细胞脂质提取物中都有降低。原子力显微镜的力谱模式也用于研究质膜制剂和全细胞脂质提取物。力测量表明,从 SL 缺乏的 LY-B 细胞获得的样品需要较低的穿透力,才能穿透比对照细胞获得的样品。质谱分析也是了解 LY-B 细胞脂质代谢重排的有用工具。LY-B 细胞中最丰富的 SL 是鞘磷脂,由于培养基中 SL 的限制,鞘磷脂减少了约 85%,生物活性脂质神经酰胺和神经节苷脂前体半乳糖脑苷脂也类似减少,胆固醇也随之减少。定量 SL 分析表明,LY-B 细胞中 SL 供应量减少 250 倍,仅导致膜 SL 减少 6 倍,这突出了这些细胞对组成变化的抵抗力。质膜组成也表现出与全细胞类似的变化,至少在质上是相似的,随着 SL 限制。在质膜中,鞘磷脂浓度与 Laurdan 荧光测量的脂质有序性之间存在线性相关性,与原子力显微镜评估的膜穿透力之间也存在线性相关性。在 SL 限制条件下,甘油磷脂也发生了较小但显著的变化。