On lipid droplets in renal interstitial cells

Summary By examining Epon sections of the renal papilla, it is demonstrated that a brief period of salt repletion in salt-depleted rats induces a considerable increase in the number of lipid droplets in the renal interstitial cells. It is supposed that this difference indicates that the lipid droplets contain substances which are physiologically important for the kidneys. Possibly, these substances are prostaglandins, which have been demonstrated previously by other authors in extracts from the renal medulla.This work was supported by a grant from the Danish State Research Foundation.The author wishes to acknowledge the support and inspiration given by E. Bojesen, M.D., and P. Leyssac, M.D., the Institute of Experimental Medicine.     

On lipid droplets in renal interstitial cells

Summary By ultracentrifugation of homogenates of rat renal papillae, a low density layer composed of small primary fluorescing lipid droplets was isolated. Probably the isolated lipid material originates mainly from the primary fluorescing lipid droplets of the renal interstitial cells. The isolated lipid droplets were mainly triglycerides, cholesterol esters and free longchain fatty acids. The long-chain fatty acid composition of the main components differed markedly from that of the analogous components of plasma and depot fat. The findings suggest that the complex lipids of the isolated lipid droplets are synthetized by the renal papilla and probably by the renal interstitial cells. The prostaglandin precursor arachidonic acid constitutes a significant fraction of the triglyceride long-chain fatty acids. Usually also small quantities of prostaglandins were present. Altogether the results suggest that the lipid droplets of the renal interstitial cells may function as a storage site for prostaglandin precursor.This work was supported by a grant from the Danish State Research Foundation. — The authors wish to acknowledge the support and inspiration given by E. Bojesen, M.D., Ph.D., the Institute of Experimental Medicine, Division of Endocrinology. — The authors wish to thank Dr. John E. Pike of the Upjohn Company, Kalamazoo, Michigan, for supplying the Prostaglandins used in this study.     

On lipid droplets in renal interstitial cells

Summary As the result of histochemical studies, it has been shown that the lipid droplets in the interstitial cells of the renal medulla of the rat contain simple saturated and unsaturated lipids. A possible correlation is suggested between the lipid droplets and the biologically active substances of a lipid character (vasodepressor lipid, medullin, and prostaglandin) which have been isolated from the renal medulla during recent years.This work was supported by a grant from the Danish State Research Foundation.     

Adipose triglyceride lipase protects renal cell endocytosis in a Drosophila dietary model of chronic kidney disease

Obesity-related renal lipotoxicity and chronic kidney disease (CKD) are prevalent pathologies with complex aetiologies. One hallmark of renal lipotoxicity is the ectopic accumulation of lipid droplets in kidney podocytes and in proximal tubule cells. Renal lipid droplets are observed in human CKD patients and in high-fat diet (HFD) rodent models, but their precise role remains unclear. Here, we establish a HFD model in Drosophila that recapitulates renal lipid droplets and several other aspects of mammalian CKD. Cell type–specific genetic manipulations show that lipid can overflow from adipose tissue and is taken up by renal cells called nephrocytes. A HFD drives nephrocyte lipid uptake via the multiligand receptor Cubilin (Cubn), leading to the ectopic accumulation of lipid droplets. These nephrocyte lipid droplets correlate with endoplasmic reticulum (ER) and mitochondrial deficits, as well as with impaired macromolecular endocytosis, a key conserved function of renal cells. Nephrocyte knockdown of diglyceride acyltransferase 1 (DGAT1), overexpression of adipose triglyceride lipase (ATGL), and epistasis tests together reveal that fatty acid flux through the lipid droplet triglyceride compartment protects the ER, mitochondria, and endocytosis of renal cells. Strikingly, boosting nephrocyte expression of the lipid droplet resident enzyme ATGL is sufficient to rescue HFD-induced defects in renal endocytosis. Moreover, endocytic rescue requires a conserved mitochondrial regulator, peroxisome proliferator-activated receptor-gamma coactivator 1α (PGC1α). This study demonstrates that lipid droplet lipolysis counteracts the harmful effects of a HFD via a mitochondrial pathway that protects renal endocytosis. It also provides a genetic strategy for determining whether lipid droplets in different biological contexts function primarily to release beneficial or to sequester toxic lipids.

A high-fat diet model of chronic kidney disease in Drosophila reveals that boosting triglyceride lipolysis in renal cells is sufficient to rescue renal cell function via a pathway involving PGC1 alpha and mitochondria.     

Lipid droplet de novo formation and fission are linked to the cell cycle in fission yeast

Cells sequester neutral lipids in bodies called lipid droplets. Thus, the formation and breakdown of the droplets are important for cellular metabolism; unfortunately, these processes are difficult to quantify. Here, we used time-lapse confocal microscopy to track the formation, movement and size changes of lipid droplets throughout the cell cycle in fission yeast Schizosaccharomyces pombe. In theory, the number of lipid droplets in these cells must increase for daughter cells to have the same number of droplets as the parent at a reference point in the cell cycle. We observed stable droplet formation events in G2 phase that were divided evenly between de novo formation of nascent droplets and fission of preexisting droplets. The observations that lipid droplet number is linked to the cell cycle and that droplets can form via fission were both new discoveries. Thus, we scrutinized each fission event for multiple signatures to eliminate possible artifacts from our microscopy. We augmented our time-lapse confocal microscopy with electron microscopy, which showed lipid droplet 'intermediates': droplets shaped like dumbbells that are potentially in transition states between two spherical droplets. Using these complementary microscopy techniques and also dynamic simulations, we show that lipid droplets can form by fission.     

Aspect ultrastructural de la lipogénèse dans le tissu adipeux brun

Summary In interscapular brown fat of the rat, appropriately processed so as to maintain membrane structures intact, lipid droplets are in fact liposomes, i.e., lipid filled vacuoles surrounded by a membrane that is related to the smooth endoplasmic reticulum. Thus the endoplasmic reticulum appears as an important component of brown fat cells. — After complete lipid depletion has been achieved by 7 days of fasting, feeding with glucose results in sudden and very conspicuous increase of smooth endoplasmic reticulum; dilatations of the perinuclear cistern and pinocytotic activity at the periphery of the cell contribute to this increase. Simultaneously the cytoplasmic matrix is heavily loaded with glycogen in particulate form. Lipogenesis, as far as can be appreciated by different degrees in electron density, takes place inside the vesicles of the endoplasmic reticulum; increase of such small lipid vacuoles then leads to reconstitution of liposomes; return to the normal aspect is completed 12 hours after the beginning of feeding. — During the phase of glycogen overloading and resulting lipogenesis, glycogen particles may be found in intercellular and pericapillary spaces; the significance of this finding is discussed. — The fundamental difference between both types of fat cells seems to be concerned with the site of lipogenesis: this takes place in the cytoplasmic matrix of white fat cells, so that lipid droplets aggregate without any limiting membrane, whereas in brown fat cells lipogenesis occurs inside cavities of the endoplasmic reticulum and lipids remain permanently enclosed in membranes. This process appears similar to what may be observed occasionally in liver and normally in adrenal cortex, and this might presumably lead to a physico-chemical understanding of the particular aspect of lipogenesis in brown fat, as compared to that in common white fat.

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Travail dédié au Professeur W. Bargmann, en témoignage d'admiration, à l'occasion de son 60e anniversaire.     

Absorption of dietary triacylglycerol by lipolysis and lipid resynthesis in the mesenteron of larvalAeshna cyanea (insecta,odonata)

Summary Voluntary uptake of triolein, margarine, and lipid-rich natural food (Tubifex) by fasting dragonfly larvae (Aeshna cyanea) led to heavy accumulations of lipid absorption droplets in the enterocytes within 2 days, while subsequent lipid clearance of the midgut epithelium took several weeks depending on the ingested lipid load. No endocytotic lipid uptake was observed after application of a molecular-dispersed fat dye. The smallest lipid droplets first appeared in the subapical groundplasm of the enterocytes and showed a reversible increase in size on their way towards the base. Lipid droplets were also observed at appropriate intervals after oral administration of oleic acid, after feeding margarine in the cold, and after injection of triolein into the isolated midgut.Comparative biochemical analysis after triolein feeding evidenced release of lipase and hydrolytic liberation of FA from TG in the midgut lumen, as well as time-dependent accumulations of TG in the midgut epithelium and of DG in the hemolymph.Oral injection of [¹⁴C] oleic acid was followed by its rapid absorption into the midgut epithelium, where it was utilized for the synthesis of MG and esterification to DG and TG. Discharge of radioactive lipid into the hemolymph occurred in the form of FA and DG, while the rectal fat body showed approximately equal labeling of the FA, DG, and TG fractions.Abbreviations AG acylglycerol - DG diacylglycerol - ER endoplasmic reticulum - FA fatty acid - MG monoacylglycerol - TG triacylglycerol Dedicated to Prof. Dr. Dr. R.Lehmensick, Bonn, in honor of his 85^th birthday.     

Ultrastructural changes in the mitochondria of the acid gland of Nasonia vitripennis (Walker) (Pteromalidae: Hymenoptera) induced by starvation

Summary The formation of mitochondrial-cytoplasmic complexes and their transformation into lipid droplets in the acid gland of Nasonia vitripennis is described. Electron microscopy and histochemistry show that lipid droplets are absent from acid glands in newly emerged, fed and re-fed insects. The droplets develop in the cytoplasm after varying periods of starvation and are not associated with acid phosphatase activity.The mature lipid droplets are rarely associated with intact mitochondria and are probably the residual end-product of the mitochondrial-cytoplasmic associations. The possible role of the associations in the maintenance of mitochondrial function and structure is discussed.We are indebted to Professor E. W. Knight-Jones in whose Department the work was carried out and to the Scientific Research Council for financial assistance to one of us (N.A.R.).     

Triglyceride depletion in THP-1 cells alters cholesteryl ester physical state and cholesterol efflux

To study macrophage lipid droplet composition and the effects of TG on cholesteryl ester (CE) physical state, hydrolysis, and cholesterol efflux, a technique was developed to remove the majority of accumulated TG with minimal effect on CE content. THP-1 macrophages were incubated with acetylated LDL, and the accumulated TG was depleted by incubation with the acyl-CoA synthetase inhibitor triacsin D in the presence of albumin. Before TG removal, all cellular lipid droplets were isotropic as determined by polarizing light microscopy. When the TG concentration was reduced, anisotropic lipid droplets were visible, indicating a change in physical state, and suggesting that TG and CE originally accumulated in mixed lipid droplets. This change in physical state of lipid droplets was associated with slower rates of CE hydrolysis and cholesterol efflux. Although lipid droplets within the same cell had a similar physical state after TG depletion, there was considerable variability among cells in the physical state of their lipid droplets.In conclusion, THP-1 macrophages store accumulated CE and TG in mixed droplets, and the proportion of CE to TG varies among cells. Reducing accumulated TG altered CE physical state, which in turn affected hydrolysis of CE and cholesterol efflux.     

Hormonal control of leydig cell differentiation

Summary The fine structure of the testicular interstitial cells of the 9-day-old mouse submitted to stimulation with human chorionic gonadotropin (HCG) is reported. As was previously described (Baillie 1964) the interstitial tissue of the prepubertal mouse testis is characterized by the presence of well differentiated epithelioid cells at a quiescent stage. They are characterized by large cytoplasmic depots of lipid droplets and glycogen particles in contrast to poorly developed membranous organelles. These cells are highly sensitive to the action of gonadotropins. Five daily injections of HCG cause their differentiation into cells with active secretory characteristics. The gonadotropin induces a marked depletion of the lipid droplets and glycogen content of the cytoplasm, concurrent with an unusual development of the membranes of the agranular endoplasmic reticulum. Golgi complexes and rough reticulum are prominent. Several changes also appear in the nucleus, especially in the nucleolus.The correlation of the present electron microscopic study of the interstitial cells under HCG stimulation with previous biochemical and physiological findings tentatively suggests that the immature Leydig cells exhibit the basic organization necessary for biosynthesis of steroid hormones.     

Work in progress correlation of renal medullary prostaglandin content and renal interstitial cell lipid droplets

Indomethacin administration and hydronephrosis in rabbits has been found to produce increases in the number and changes in the composition of the lipid droplets in the renal medullary interstitial cells. The response to indomethacin, a prostaglandin synthetase inhibitor, was dose dependent.Work is in progress to assess the effects of other non-steroidal antiinflammatory drugs on the renal inner medulla and the interstitial cell lipid droplets.     

Triglyceride containing lipid droplets and lipid droplet-associated proteins

PURPOSE OF REVIEW: Cytosolic lipid droplets are now recognized as dynamic organelles. This review summarizes our current understanding of the mechanisms involved in the formation of lipid droplets, the importance of lipid droplet-associated proteins and the link between lipid droplet accumulation and development of insulin resistance. RECENT FINDINGS: Lipid droplets are formed as primordial droplets and they increase in size by fusion. This fusion process requires the alpha-soluble N-ethylmaleimide-sensitive factor adaptor protein receptor SNAP23, which is also involved in the insulin-dependent translocation of a glucose transporter to the plasma membrane. Recent data suggest that SNAP23 is the link between increased lipid droplet accumulation and development of insulin resistance. Lipid droplets also form tight interactions with other organelles. Furthermore, additional lipid droplet-associated proteins have been identified and shown to play a role in droplet assembly and turnover, and in sorting and trafficking events. SUMMARY: Recent studies have identified a number of key proteins that are involved in the formation and turnover of lipid droplets, and SNAP23 has been identified as a link between accumulation of lipid droplets and development of insulin resistance. Further understanding of lipid droplet biology could indicate potential therapeutic targets to prevent accumulation of lipid droplets and associated complications.     

Increased lipid droplet accumulation associated with a peripheral sensory neuropathy

Hereditary sensory neuropathy type 1 (HSN-1) is an autosomal dominant neurodegenerative disease caused by missense mutations in the SPTLC1 gene. The SPTLC1 protein is part of the SPT enzyme which is a ubiquitously expressed, critical and thus highly regulated endoplasmic reticulum bound membrane enzyme that maintains sphingolipid concentrations and thus contributes to lipid metabolism, signalling, and membrane structural functions. Lipid droplets are dynamic organelles containing sphingolipids and membrane bound proteins surrounding a core of neutral lipids, and thus mediate the intracellular transport of these specific molecules. Current literature suggests that there are increased numbers of lipid droplets and alterations of lipid metabolism in a variety of other autosomal dominant neurodegenerative diseases, including Alzheimer’s and Parkinson’s disease. This study establishes for the first time, a significant increase in the presence of lipid droplets in HSN-1 patient-derived lymphoblasts, indicating a potential connection between lipid droplets and the pathomechanism of HSN-1. However, the expression of adipophilin (ADFP), which has been implicated in the regulation of lipid metabolism, was not altered in lipid droplets from the HSN-1 patient-derived lymphoblasts. This appears to be the first report of increased lipid body accumulation in a peripheral neuropathy, suggesting a fundamental molecular linkage between a number of neurodegenerative diseases.     

The interstitial cells in the renal medulla of rat,rabbit, and gerbil in different states of diuresis

Summary The inner zone of the renal medulla of rats, gerbils, and rabbits was investigated to determine whether or not there are any characteristic ultrastructural differences between the interstitial cells of these species. The effects on the interstitial cells of water deprivation and water loading were also investigated.In all three species, the Type 1 interstitial cells, the lipid containing cells, were abundant and their distribution and topographical relations as well as their general ultrastructure were similar. The previously reported significantly higher frequency in desert rats could not be confirmed. Although the lipid droplets of the interstitial cells were smaller in gerbils and rabbits when compared to rats, their fine structure was similar. Their electron dense outer zone was sometimes associated with a granular material and/or a lamellar material with a periodicity of about 40 Å resembling phospholipid myelin figures.Water-loaded rats showed a considerable increase in the number of lipid droplets when compared to dehydrated or untreated animals. In contrast, the interstitial cells of waterloaded gerbils and rabbits were depleted of lipid droplets.We are indebted to Professor A.B. Maunsbach for valuable discussions and criticism and to Mrss. Hanne Weiling and Birthe Overgaard for competent technical assistance. The gerbils used in this study were a gift from Leo AB, Helsingborg, Sweden. This study was supported by the Danish Medical Research Council (J. nos. 512-1067, 512-1545, 512-3633)     

Free fatty acids,lipid peroxidation,and lysosomal enzymes in experimental focal cerebral ischemia in primates: Loss of lysosomal latency by lipid peroxidation

Experimental focal cerebral ischemia was produced in monkeys (Macaca radiata) by occlusion of the right middle cerebral artery (MCA). The release of the lysosomal glycosidases, -d-hexosaminidase, -l-fucosidase and -d-mannosidase into the soluble fraction in the right basal ganglia of the experimental animals was measured at different periods from 30 min to 12 hr after occlusion and compared with the corresponding sham operated control animals. There was a significant increase in the released lysosomal enzymes in the MCA occluded animals at all periods and particularly at 4 hr after occlusion. The CSF from the experimental animals also showed elevated levels of hexosaminidase and fucosidase. The free fatty acids (FFA) measured in the basal ganglia at 30 min and 2 hr after occlusion showed a 100 fold increase in the experimental animals. The predominant fatty acid released was linoleic acid (18:2) followed by arachidonic acid (20:4). Lipid peroxidation in the basal ganglia measured by the thiobarbituric acid (TBA) reaction in the presence or absence of ascorbic acid also showed a significant increase in the experimental animals at all periods with a maximum at 30 min to 2 hr after occlusion. In order to assess whether lipid peroxidation causes damage to the lysosomes and release of the enzymes, a lysosome enriched P₂ fraction from the normal monkey basal ganglia was prepared and the effect of peroxidation studied. Maximum peroxidation in the P₂ fraction was observed in the presence of arachidonic acid, ascorbic acid and Fe²⁺. There was a good correlation between the extent of lipid peroxidation and the in vitro release of lysosomal hexosaminidase from the P₂ fraction. Anti-oxidants which strongly inhibited lipid peroxidation in the P₂ fraction prevented the release of hexosaminidase. The results suggested that in ischemia produced by MCA occlusion lipid peroxidation which damages the lysosomal membrane causes the release of lysosomal hydrolytic enzymes.Abbreviations used BHA butylated hydroxyanisole - BHT butylated hydroxytoluene - FFA free fatty acids - MCA middle cerebral artery - MDA malonaldehyde - PUFA polyunsaturated fatty acids - TBA thiobarbituric acid     

Biomodulator-mediated susceptibility of endogenous lipid droplets from rat adipocytes to hormone-sensitive lipase

The amount of fatty acid release by a fat cell homogenate without pretreatment with epinephrine was found to be slightly more than that released from fat cells by epinephrine, suggesting that fat cells contain high lipolytic activity even in the absence of lipolytic agents. Fat cells contain high hormone-sensitive lipase activity (1383 mumole free fatty acids/g/hr) in the absence of epinephrine, and addition of epinephrine to the cells did not increase the activity, significantly. Like epinephrine, DBcAMP and/or theophylline also elicited marked release of glycerol from fat cells without activating the hormone-sensitive lipase activity. However, although fat cells contain a large amount of hormone-sensitive lipase, lipolysis was negligible in the absence of these lipolytic agents. These results suggest that lipolytic agents such as epinephrine, DBcAMP, and theophylline induce lipolysis in fat cells through some mechanism other than activation of hormone-sensitive lipase and that in the absence of lipolytic agents, some system in fat cells inhibits lipolysis of endogenous lipid droplets by hormone-sensitive lipase. The lipid droplets in fat cells consist mainly of triglyceride with phospholipids, cholesterol, carbohydrate, and protein as minor constituents. The phospholipid fraction was found to consist of 75% phosphatidylcholine and 25% phosphatidylethanolamine. Of the minor constituents of endogenous lipid droplets, only phosphatidylcholine strongly inhibited hormone-sensitive lipase activity in a [3H]triolein emulsion. These results suggest that phosphatidylcholine in endogenous lipid droplets may be responsible for inhibition of hormone-sensitive lipase. Then, a cell-free system was established in which epinephrine, DBcAMP, and theophylline stimulated lipolysis of endogenous lipid droplets from fat cells by lipase solution. In this system, these lipolytic agents did not induce lipolysis in the absence of added lipase. Lipolysis in the mixture of the endogenous lipid droplets and lipase solution was accelerated by phospholipase C with concomitant loss of epinephrine-induced lipolysis. After pretreatment of the endogenous lipid droplets with phospholipase C, these lipolytic agents no longer induced lipolysis. Pretreatment of the endogenous lipid droplets with phospholipase C reduced their phospholipid content with the formation of phosphorylcholine, but did not affect their triglyceride and cholesterol contents. Treatment of the endogenous lipid droplets with phospholipase D did not affect lipolysis in the cell-free system. These results suggest that phosphatidylcholine in the endogenous lipid droplets may inhibit their lipolysis by hormone-sensitive lipase in fat cells and also be involved in the mechanisms of the stimulatory effects of epinephrine, DBcAMP, and theophylline on lipolysis.     

Sphingosine 1-phosphate metabolism and insulin signaling

Insulin is the main anabolic hormone secreted by β-cells of the pancreas stimulating the assimilation and storage of glucose in muscle and fat cells. It modulates the postprandial balance of carbohydrates, lipids and proteins via enhancing lipogenesis, glycogen and protein synthesis and suppressing glucose generation and its release from the liver. Resistance to insulin is a severe metabolic disorder related to a diminished response of peripheral tissues to the insulin action and signaling. This leads to a disturbed glucose homeostasis that precedes the onset of type 2 diabetes (T2D), a disease reaching epidemic proportions. A large number of studies reported an association between elevated circulating fatty acids and the development of insulin resistance. The increased fatty acid lipid flux results in the accumulation of lipid droplets in a variety of tissues. However, lipid intermediates such as diacylglycerols and ceramides are also formed in response to elevated fatty acid levels. These bioactive lipids have been associated with the pathogenesis of insulin resistance. More recently, sphingosine 1-phosphate (S1P), another bioactive sphingolipid derivative, has also been shown to increase in T2D and obesity. Although many studies propose a protective role of S1P metabolism on insulin signaling in peripheral tissues, other studies suggest a causal role of S1P on insulin resistance. In this review, we critically summarize the current state of knowledge of S1P metabolism and its modulating role on insulin resistance. A particular emphasis is placed on S1P and insulin signaling in hepatocytes, skeletal muscle cells, adipocytes and pancreatic β-cells. In particular, modulation of receptors and enzymes that regulate S1P metabolism can be considered as a new therapeutic option for the treatment of insulin resistance and T2D.     

The fine structure of the mouse adrenal X zone

Summary Electron microscopically the adrenal X zone was examined in the fourteen SMA female mice aged 40 and 70 days. At these ages, the X zone showed no signs of degeneration. The X zone cell was somewhat smaller than the permanent cortical cell.The mitochondria in the X zone cell were quite bizarre in shape, provided with tubules or cristae. Many intramitochondrial bodies very similar to the cytoplasmic lipid droplets were found in the X zone. A few lipid droplets and globules were also noticed in this zone. The lipid droplets may possibly be formed within the mitochondria.The light and dark cells were differentiated. For the light cells, scant mitochondria and tubular granular endoplasmic reticulum were characteristic in contrast to the abundant mitochondria and multi-lamellated agranular endoplasmic reticulum in the dark cells. The cellular variety in density was discussed with regard to steroid synthesis.The author wishes to express his sincere appreciation to Prof. H. Tauchi, The 2nd Department of Pathology, Nagoya University School of Medicine, for kind advice, to Dr. M. Hoshino for helpful suggestion, and to Mr. J. Aoki for excellent technical assistance.     

Physiological status and change of cytoplasmic lipid droplets in the pheromone-producing cells of the silkmoth, Bombyx mori (Lepidoptera, Bombycidae)

Changes in size and number of cytoplasmic lipid droplets were quantified in the pheromone gland (PG) of Bombyx mori before and after adult eclosion. Two days before eclosion, size and number of droplets are small (diameter is 2-7 microm) and few. The formation and significant proliferation of larger droplets (5-12 microm) take place between 2 days and 1 day before eclosion. From the day of emergence until day 3 a fluctuation in size and number of lipid droplets during the photophase (4h intervals) is observed. The changes are more characteristic and dramatic on the day of emergence and first day, while attenuation of these changes can be observed from the second day and seems to disappear by day 4. Bombykol content, at each respective time, is in good correlation with the observed fluctuation in lipid droplet parameters. Highest bombykol production daily is observed towards the early evening, when lipid droplets are the smallest (2-4 microm) and most numerous. By day 4, however, this regularity also ceases. In 24h old mated females PG cell structure is quite similar to newly emerged ones. In glands of 72 h old decapitated females the formation of 'extra' large lipid droplets is remarkable. In vivo pheromone biosynthesis activating neuropeptide (PBAN) treatment, however, induced the formation of many small droplets, although numerous larger ones also remained. The morphological changes in lipid droplets and cellular dynamics associated with the external signal of PBAN in the PG suggest a storage-pool function of the lipid droplets.