Biosynthesis of galactolipids by enzyme preparations from spinach leaves

The pH optimum for galactolipid synthesis from UDP-galactose by spinach chloroplasts is 7.2 in Tris-HCl or phosphate buffer. The products include sterol glycosides, trigalactosyl diglyceride (tentatively identified), digalactosyl diglyceride, and monogalactosyl diglyceride in increasing order of quantity. The proportion of monogalactosyl diglyceride decreases and that of digalactosyl diglyceride increases as the pH is lowered. The galactolipid synthesis is quite resistant to elevated temperature; maximal incorporation of galactose from UDP-galactose was observed at 45 degrees C. The proportion of monogalactosyl diglyceride was greater at the higher temperatures. As much as 40% of the galactolipid-synthesizing capability of a spinach leaf homogenate is not sedimented by centrifugation for 60 min at 100,000 g. An acetone powder of spinach chloroplasts contains enzymes which catalyze galactolipid synthesis. This preparation is dependent on added diglycerides in order to make galactolipid, whereas the chloroplast preparation is not dependent on added diglycerides. Molecular species of diglycerides were compared as requirements for galactolipid synthesis. The requirement was satisfied best by the diglycerides of highest unsaturation. Methylation of the free hydroxyl of the diglyceride eliminated the effectiveness.     

BIOSYNTHESIS AND COMPOSITION OF BRAIN GALACTOLIPIDS IN NORMAL AND HYPOTHYROID RATS

Abstract— Newborn rats were rendered hypothyroid by methimazole treatment. Incorporation of [1-¹⁴C]galactose both in vivo and in vitro into brain cerebrosides of hypothyroid rats was significantly lower than in normals. Biosynthesis of sulphatides was affected by hypothyroidism to a smaller extent than cerebrosides. Assay of cerebroside biosynthesis from [1-¹⁴C]galactose or UDP-[1-¹⁴C]galactose by brain preparations revealed that incorporation of the sugar in both cases is affected to the same extent by methimazole treatment, suggesting that the phenomenon is not due to impairment of the nucleotide biosynthesis. A radioactive galactolipid tentatively characterized as glycerogalactolipid was synthesized in vitro and its biosynthesis was reduced to a large extent in the brain preparations from hypothyroid rats. The fatty acid composition of cerebrosides and sulphatides from the brains of hypothyroid rats was found to be different from that of normal rats. The percentage of normal C₂₄ fatty acids was significantly decreased in the methimazole-treated rats. Brain sphingomyelin fatty acids did not differ between normal and hypothyroid rats.     

Biosynthesis of lipids in chloroplasts isolated from jack pine needles

Suspensions of isolated pine needle chloroplasts were shown to incorporate galactose from UDP galactose-[¹⁴C] into galactolipids. The incorporation of the label among galactolipids was always considerably higher in the monogalactosyl diglycerides than in the digalactosyl diglycerides. The galactosyl incorporation into both galactolipid fractions was optimal at pH 8.0 and was inhibited by sulphydryl reagents (p-chloromercuribenzoate, N-ethyl maleimide and CdCl₂). The chloroplast preparations were also able to biosynthesize various phospholipids and galactolipids from palmitoyl-[1-¹⁴C]-CoA; the major portion of the label appeared in phosphatidyl choline. The incorporation of palmitic-[1-¹⁴C] acid into various lipids was very poor compared to that of palmitoyl-[1-¹⁴C]-CoA. However, addition of ATP and CoA markedly stimulated lipid biosynthesis from palmitic-[1-¹⁴C] acid, suggesting the presence of activating enzymes. These chloroplast suspensions did not show any de novo fatty acid synthesis.     

Quantitative and compositional changes in monogalactosyl and digalactosyl diglycerides during light-induced formation of chloroplasts in Euglena gracilis

The formation of chloroplasts in dark-grown cells of Euglena gracilis was induced by exposing the cells to constant illumination. Following a lag, the cells accumulated chlorophyll and galactosyl diglycerides simultaneously at almost linear rates. The monogalactosyl diglyceride content rose from approximately 2 micromoles in 100 mg of dark-grown cells to 27 micromoles in fully green cells; the digalactosyl diglyceride content increased from 1 micromole to 11 micromoles. The digalacto compounds increased more rapidly than the monogalacto compounds at first, but their rate of accumulation began to diminish long before greening of the cell was complete. The sole exception was the digalactosyl diglyceride fraction that contained hexadecadienoic (16:2) fatty acid. This fraction increased continuously during greening. As accumulation of the digalacto compounds diminished, that of the monogalacto compounds increased. Towards the end of greening, the major fatty acids were 16:2, 16:3, 16:4, 18:2, and 18:3 in the monogalacto and 16:2 in the digalacto compounds. The results of this study suggest that monogalactosyl and digalactosyl diglycerides that contain particular fatty acid components have a function in the assembly of chloroplasts.     

The fatty acid composition of fern lipids

Fatty acid compositions of the lipids isolated from the pinnae of four fern species show that they differ from those of the lipids in leaves of higher plants in having C-20 polyunsaturated acids, mainly arachidonic acid. As in higher plants, the ω-3 polyunsaturated acids are concentrated in the monogalactosyl diglycerides. Variations are found both in the fatty acid compositions and in the monogalactosyl/digalactosyl diglyceride ratio during the growing season.     

Monogalactosyl and digalactosyl diglycerides from heterotrophic, hetero-autotrophic, and photobiotic Euglena gracilis

The lipid of Euglena gracilis, dark-grown in a complete medium, contained 2% galactose. The lipid of Euglena gracilis, light-grown in either a complete or an inorganic medium, contained 13-14% galactose. Pure monogalactosyl and digalactosyl diglyceride fractions, isolated by column plus thin-layer chromatography, contained 50% of the lipid-bound galactose of dark-grown cells, and 80% of that of light-grown cells. Molar ratios of monogalactosyl to digalactosyl compounds ranged from 2 to 3. The results show that galactosyl diglycerides, stored in large amount in light-grown cells, persist in small amount in the dark-grown cells. Fatty acids in both the monogalactosyl and the digalactosyl diglycerides were mainly of the 16- and 18-carbon varieties, with high proportions of trienes. The monogalactosyl diglycerides were rich in hexadecatetraenoic acid. Strictly photobiotic cells had twice as much hexadecadienoic and hexadecatetraenoic acids in their monogalactosyl diglycerides, and three times as much hexadecadienoic and octadecadienoic acids in their digalactosyl diglycerides as did illuminated cells grown in a complete medium. Dark-grown (obligate) heterotrophs contained galactosyl diglycerides with high percentages of monoenes. Great compositional variations in the galactosyl diglycerides are thus induced by light and also by nonlipid exogenous metabolites.     

Lipid biosynthesis by chloroplasts from olive tree leaves

In vitro incubation of isolated chloroplasts from young olive tree leaves ( Olea europaea L. cv. Marteño) in acetate-1-¹⁴C showed a high labelling of saturated fatty acids (palmitic + stearic) and, above all, of the monounsatured ones (oleic); the low biosynthetic rate of α-linolenic acid being noteworthy. These fatty acids are mainly found as free ones, or incorporated in mono-and diglyceride molecules. Phosphoand galactolipids, the most abundant acyl-lipid components of chloroplast lamellae, showed low incorporation rates. The fatty acid synthesis by isolated chloroplasts depends on exogenous CoA, ATP, NADPH and, especially, on added ACP (acyl carrier protein) preparation from Escherichia coli , whereas it was strongly inhibited by Triton X-100.
In vivo experiments with acetate-1-¹⁴C infiltration into young excised leaves showed a high labelling of chloroplast phospholipids, but a low ¹⁴C incorporation into galactolipids, a remarkable feature because these latter are main components of chloroplast lamellae. The high biosynthetic rate of α-linolenic acid is noteworthy and appears mainly linked to monogalactosyldiglycerides. Also the low incorporation of saturated fatty acids to neutral lipids is remarkable. The low in vitro synthesis of α-linolenic acid in comparison with that of the in vivo conditions, suggests the existence of a cooperation between chloroplasts and other parts of the cell to carry out the synthesis of this compound.     

Polygalactolipids in Spinach Chloroplast

Two polygalactolipids, designated as components A and B, were isolated from spinach chloroplasts and were also obtained from glycolipid products synthesized with chloroplast enzymes using uridine diphosphate galactose as a galactose donor. These lipids were purified by column and thin layer chromatography. Chemical analysis of component A indicates that the lipid is trigalactosyl diglyceride, whereas component B behaves like tetragalactosyl diglyceride on a thin layer plate. The major fatty acid in trigalactosyl diglyceride was alpha-linolenic acid. Relative amount (molar ratio) of galactolipids in spinach chloroplasts was monogalactosyl diglyceride:digalactosyl diglyceride:trigalactosyl diglyceride:(tetragalactosyl diglyceride) = 60:30:5:1.     

Docosahexaenoic acid synthesis from n-3 fatty acid precursors in rat hippocampal neurons

Docosahexaenoic acid (DHA), the most abundant n-3 polyunsaturated fatty acid in the brain, has important functions in the hippocampus. To better understand essential fatty acid homeostasis in this region of the brain, we investigated the contributions of n-3 fatty acid precursors in supplying hippocampal neurons with DHA. Primary cultures of rat hippocampal neurons incorporated radiolabeled 18-, 20-, 22-, and 24-carbon n-3 fatty acid and converted some of the uptake to DHA, but the amounts produced from either [1-¹⁴C]α-linolenic or [1-¹⁴C]eicosapentaenoic acid were considerably less than the amounts incorporated when the cultures were incubated with [1-¹⁴C]22:6n-3. Most of the [1-¹⁴C]22:6n-3 uptake was incorporated into phospholipids, primarily ethanolamine phosphoglycerides. Additional studies demonstrated that the neurons converted [1-¹⁴C]linoleic acid to arachidonic acid, the main n-6 fatty acid in the brain. These findings differ from previous results indicating that cerebral and cerebellar neurons cannot convert polyunsaturated fatty acid precursors to DHA or arachidonic acid. Fatty acid compositional analysis demonstrated that the hippocampal neurons contained only 1.1–2.5 mol% DHA under the usual low-DHA culture conditions. The relatively low-DHA content suggests that some responses obtained with these cultures may not be representative of neuronal function in the brain.     

Transport in C4 mesophyll chloroplasts. Characterization of the pyruvate carrier

1. Evidence is presented for high rates of carrier-mediated uptake of pyruvate into the stroma of intact mesophyll chloroplasts of the C₄ plant Digitaria sanguinalis, but not the chloroplasts of the C₃ plant Spinacea oleracea. Uptake of pyruvate in the dark with the C₄ mesophyll chloroplasts was followed using two techniques: uptake of [¹⁴C]pyruvate as determined by silicon oil centrifugal filtration and uptake as indicated by absorbance changes at 535 nm (shrinkage/swelling) after addition of 0.1 M pyruvate salts.

2. Uptake of the pyruvate anion by an electrogenic carrier is suggested to be the major mode of transport. Chloroplast swelling was observed in potassium pyruvate plus valinomycin and uptake of [¹⁴C]pyruvate was inhibited by membrane-permeant anions. Valinomycin reduced uptake in the absence of external potassium and the inhibition could be reversed by addition of external potassium.

3. Uptake of pyruvic acid (or a pyruvate ⁻/OH⁻ antiport) is ruled unlikely since [¹⁴C]pyruvate uptake was relatively independent of the pH gradient across the envelope and addition of pyruvate to chloroplasts did not result in an alkalization of the medium. The low rate of swelling observed in ammonium pyruvate may be due to non-mediated permeation of pyruvic acid, which is possible only at high pyruvate concentrations.

4. The concentration of pyruvate in the stroma increased with external concentration over the range tested (up to 40 mM) but the concentration ratio (internal/external) was always less than one. The steady-state concentration of [¹⁴C]pyruvate in the stroma was dependent on the ionic strength of the medium, with saturation at roughly I = 0.04 M, while accumulation of the membrane-permeant cation tetraphenylmethylphosphonium decreased with increasing ionic strength. This suggests that ionic strength modifies a membrane potential (inside negative) across the envelope and that pyruvate uptake responds to the magnitude and direction of that potential (−80 mV at low ionic strength).

5. Chloride and inorganic phosphate were potent inhibitors of [¹⁴C]pyruvate uptake. Of the sulfhydryl reagents tested, N-ethylmaleimide was not inhibitory while mersalyl completely blocked [¹⁴C]pyruvate uptake and swelling in potassium pyruvate plus valinomycin. Pyruvate uptake, as measured by valinomycin induced swelling in potassium pyruvate, was highly temperature sensitive, with an energy of activation of 39 kcal/mol above 9 °C.

6. Phenylpyruvate, -ketoisovalerate, -ketoisocaproate, -cyano-4-hydroxycinnamic acid and -cyanocinnamic acid inhibited [¹⁴C]pyruvate but not [¹⁴C]-acetate uptake in the dark and also reduced pyruvate metabolism by the chloroplasts in the light.     

Interrelationships between fatty acid biosynthesis and acyl-lipid synthesis in Chlorella vulgaris

1. Fatty acid synthesis from [2-(14)C]acetate by Chlorella vulgaris cells grown and incubated in the dark is limited almost entirely to the production of saturated and monoenoic acids. 2. In light-incubated cells, both saturated and polyunsaturated fatty acids are rapidly synthesized. 3. Two groups of lipids can be distinguished in both dark- and light-incubated cells. The first group, consisting of phosphatidyl-glycerol, monogalactosyl diglyceride, lecithin and neutral glyceride, has a very high turnover rate for certain fatty acids. The second group, consisting of digalactosyl diglyceride, sulpholipid, phosphatidylethanolamine and phosphatidylinositol, has a slow turnover of fatty acids. 4. The lipids with rapid fatty acid turnover may be involved in the sequences of saturated and unsaturated fatty acid synthesis. A classification of lipids is made on the basis of their suggested functions.     

Similarities and differences in lipid metabolism of chloroplasts isolated from 18:3 and 16:3 plants

Photosynthetically active chloroplasts retaining high rates of fatty acid synthesis from [1-¹⁴C]acetate were purified from leaves of both 16:3 (Solanum nodiflorum, Chenopodium album) and 18:3 plants (Amaranthus lividus, Pisum sativum). A comparison of lipids into which newly synthesized fatty acids were incorporated revealed that, in 18:3 chloroplasts, enzymic activities catalyzing the conversion of phosphatidate to diacylglycerol and of diacylglycerol to monogalactosyl diacylglycerol (MGD) were significantly less active than in 16:3 chloroplasts. In contrast, labeling rates of MGD from UDP-[¹⁴C]gal were similar for both types of chloroplasts.

The composition and positional distribution of labeled fatty acids within the glycerides synthesized by isolated 16:3 and 18:3 chloroplasts were similar and in each case only a C18/C16 diacylglycerol backbone was synthesized. In nodiflorum chloroplasts, C18:1/C16:0 MGD assembled de novo was completely desaturated to the C18:3/C16:3 stage.

Whereas newly synthesized C18/C18 MGD could not be detected in any of these chloroplasts if incubated with [¹⁴C]acetate after isolation, chloroplasts isolated from acetate-labeled leaves contained MGD with labeled C18 fatty acids at both sn-1 and sn-2 positions. Taken together, these results provide further evidence on an organellar level for the operation of pro- and eucaryotic pathways in the biosynthesis of MGD in different groups of plants.

     

Photosynthetic CO2 fixation by chloroplast populations isolated by a polymer two-phase technique

Preparations of spinach chloroplasts, essentially free from contamination by other cellular material or whole cells, incorporated ¹⁴C almost entirely into glycolate, a polyglucan (probably starch) and intermediates of the Calvin cycle and starch synthesis. About 70% of the ¹⁴C was found in dihydroxyacetone phosphate, 3-phosphoglycerate and glycolate. Only small amounts were found in sucrose and amino acids.

On the other hand, preparations consisting of particles containing chloroplasts surrounded by a membrane-bound cytoplasmic layer including mitochondria and microbodies, gave a much broader spectrum of ¹⁴C-labelled products. Much less of the ¹⁴C was found in dihydroxyacetone phosphate and 3-phosphoglycerate. Instead, sucrose, malate, aspartate, alanine and some other amino acids contained about 40% of the ¹⁴C incorporated. It is concluded that sucrose is synthesized by cooperation between the chloroplast and the surrounding layer.     

Mechanism of chain initiation by dextransucrase: Absence of terminal ‘sucrose’ linkage in newly synthesized dextran chains

Dextran was synthesized using dextransucrase from Streptococus sanguis 10558 and (F)-[¹⁴C]sucrose as substrate to test the possibility that sucrose may be the initial acceptor for glucose. If sucrose is the initial acceptor, then dextran chains should have [¹⁴C] fructose in a terminal ‘sucrose’ linkage which can be cleaved under mild conditions. Although incorporation of [¹⁴C]fructose into dextran was observed, the label was not released by mild hydrolysis, indicating that sucrose is not the initiator for dextran synthesis. Incorporation of [¹⁴C]fructose into dextran might represent its ability to act as an acceptor, as suggested by the isolation of leucrose as a by-product in the reaction.     

Degradation of monogalactosyl diglyceride and diagalactosyl diglyceride by sheep pancreatic enzymes

1. Saline extract of sheep pancreas acetone-dried powder was shown to catalyse acyl ester hydrolysis of spinach leaf galactosyl diglycerides and also galactosylglucosyl diglyceride of Lactobacillus casei. 2. Sodium deoxycholate stimulated the enzyme activity. Ca(2+) had no effect on the hydrolysis of monogalactosyl diglyceride, but it enhanced that of digalactosyl diglyceride. When added together, there was considerably less activity with both the substrates. 3. Optimal hydrolysis was observed at pH7.2. 4. The initial point of hydrolysis was at position-1, leading to the formation of monogalactosyl monoglyceride and digalactosyl monoglyceride. Further hydrolysis to the corresponding galactosylglycerols and later to galactose and glycerol was also observed, indicating the presence of alpha- and beta-galactosidases in the enzyme preparation. 5. Formation of monogalactosyl diglyceride from digalactosyl diglyceride by the action of alpha-galactosidase was noted. 6. Monogalactosyl diglyceride was also hydrolysed by beta-galactosidase to a limited extent, giving rise to diacylglycerol and galactose. 7. Attempts at purification of monogalactosyl diglyceride acyl hydrolase by using protamine sulphate treatment, Sephadex G-100 filtration and DEAE-cellulose chromatography gave a partially purified enzyme which showed 9- and 81-fold higher specific activity towards monogalactosyl diglyceride and digalactosyl diglyceride respectively. This still showed acyl ester hydrolysis activity towards methyl oleate, phosphatidylcholine and triacylglycerol. 8. When sheep, rat and guinea-pig tissues were compared, guinea-pig tissues showed the highest activity towards both monogalactosyl diglyceride and digalactosyl diglyceride. In all the species pancreas showed higher activity than intestine.     

Bradykinin effects on phospholipid metabolism and its relation to arachidonic acid turnovers in neuroblastoma × glioma hybrid cells (NG 108-15)

In neuroblastoma × glioma hybrid cells (NG 108-15) labelled with [³²P]-trisodium phosphate, [³H]-inositol and [¹⁴C]-arachidonic acid, bradykinin stimulated the hydrolysis of phosphatidylinositol 4,5-bisphosphate (PIP₂) while it had no effect on the release of [¹⁴C]-arachidonic acid (AA). The effect on PIP, was time- and dose-dependent with a maximal effect on [³H]-inositol- and [³²P]-labelled cells after 10–30 s of stimulation with 10⁻⁶ M bradykinin. However, the hydrolysis of [¹⁴C]-AA labelled PIP₂ was delayed compared to the effect on [³H]- and [¹⁴C]-PIP₂ and was not detectable until after 60 s of stimulation. Bradykinin stimulation resulted in an increased formation of [³H]-inositol phosphates (IP) and [³²P]- and [¹⁴P]- and [¹⁴C]-phosphatidic acid (PA) but the time course for PA formation did not allow the time-course for PIP₂ hydrolysis. A reduced labelling of [²³P]- and [¹⁴C]-phosphatidylcholine was also found in stimulated cells suggesting that PA may derive from other sources than PIP₂. In conclusion, our results indicate that bradykinin activates phospholipase C, but not phospholipase A₂, in NG 108-15 cells.     

Quantitative aspects of triacylglycerol metabolism and sterol synthesis from 14C-acetate in etiolated seedlings of Euphorbia lambii

Triacylglycerols occur in both the endosperm and embryo of Euphorbia lambii seeds. Upon germination, the amount of these neutral lipids in the endosperm decreased with 1.06 mg fatty acid day^-1. The embryo contained 1.4 mg fatty acids in the triacylglycerols and this value declined slowly to 0.4 mg seedling^-1 during the 8 day period of endosperm depletion. Radioactive acetate was rapidly taken up by the cotyledons of intact seedlings, translocated throughout the entire seedling, and up to 10.5% of the ¹⁴C proceeded to the sterols and latex triterpenols. Maximum uptake values of 1.4 μmol seedling^-1 day^-1 of acetate were measured. Acetate uptake and subsequent incorporation into sterols and triterpenols decreased substantially in the presence of increasing amounts of sucrose (up to 0.3 M). Traces of acetate did not effect [¹⁴C]-sucrose uptake and corresponding synthesis of [¹⁴C]-sterols and triterpenols, but increased concentrations of acetate (0.05 M and up) reduced both uptake of sucrose and its conversion into unsaponifiable lipids.
The uptake capacity of the cotyledons for [¹⁴C]-glycerol exceeded the daily production in the endosperm, but only a small amount of label proceeded to the sterols and triterpenols. [¹⁴C]-Triacylglycerols were never detected in the seedling, regardless of the labeled substrate used. Although acetate is an efficient precursor in triterpenol and sterol synthesis, the uptake capacity of the cotyledons for this metabolite is too small in relation to the daily production of water soluble substrates in the endosperm. If acetate is released by the endosperm, only a marginal contribution towards triterpenol and sterol synthesis in the seedling is to be anticipated from this substrate.     

Lipid synthesis by oligodendrocytes from adult pig brain maintained in long-term culture

Oligodendrocytes were isolated from adult pig brain and cultivated for 18–24 days. [¹⁴C]acetate, [³H]galactose or [³⁵S]sulfate were added to the medium for an additional 24 h. Lipids were extracted and separated by high-performance thin-layer chromatography. The labeled lipids were studied by fluorography and scintillation counting. [¹⁴C]acetate was incorporated in decreasing order into neutral lipids, phosphatidylcholine, ethanolamine phosphatides, galactocerebrosides, phosphatidylinositol, phosphatidylserine, sulfatides and sphingomyelin. From the [¹⁴C]acetate incorporated into ethanolamine and choline phosphatides, 71.6 and 14.8%, respectively, were found in plasmalogens. Among neutral lipids, [¹⁴C]acetate labeled not only cholesterol but also large amounts of triglycerides. No cholesterol esters were synthesized. [³H]galactose primarily labeled galactocerebrosides, sulfatides, and monogalactosyl diglyceride. [³⁵S]sulfate incorporation was restricted to sulfatides. Together with our previous results concerning proteins, these data show that: (1) oligodendrocytes remain highly differentiated in long-term cultures; (2) they are able to synthesize the major components of myelin; (3) they synthesize surprisingly high amounts of triglycerides and of monogalactosyl diglyceride, a marker for myelination.     

Activation of [C]chlorobiphenyls to protein-binding metabolites by rat liver microsomes

The irreversible binding of [¹⁴C]2,2′-di- and [¹⁴C]2,4,5,2′,4′,5′-hexachlorobiphenyl ([¹⁴C]DCB and [¹⁴C]HCB) to protein was studied in the presence of rat liver microsomes and a NADPH-generating system. Protein-bound radioactivity was found with [¹⁴C]DCB but not with [¹⁴C]HCB. The binding of ¹⁴C-metabolites was increased by pretreatment of the rats with phenobarbital or polychlorinated biphenyls. Protein binding was linear for 80 min. In contrast, monohydroxy-metabolites of DCB were formed and degraded within 40 min. Inhibition of secondary oxidation of DCB by scavening superoxide anions or by glucuronidation of the monophenols markedly decreased the protein binding. Addition of trichloropropene oxide or styrene oxide, both inhibitors of epoxide hydrase, did not significantly stimulate the binding. The results suggest that the majority of reactive metabolites of DCB arise from secondary metabolism, i.e., the subsequent oxidation of the phenolic metabolites. Arene oxides, the primary products, appear to play a minor role in the protein binding of DCB.     

Photodecomposition of Monogalactosyl Diglyceride Mediated by Chlorophyll

In acetone extracts the presence of chlorophyll a and b caused enhanced photodecomposition of monogalactosyl diglyceride. Irradiation of isolated etioplasts containing chlorophyllide and monogalactosyl diglyceride caused photodecomposition of chlorophyllide but not of monogalactosyl diglyceride. In irradiated etiochloroplasts containing chlorophyll a and monogalactosyl diglyceride both components were photodecomposed. During photodecomposition of monogalactosyl diglyceride the fatty acid composition remained constant.