Saving for the future: Pre‐winter uptake of algal lipids supports copepod egg production in spring

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Biosynthesis of carotenoids in plastids of plants

This review deals with various aspects of the biosynthesis of carotenoids in chromoplasts and chloroplasts of green algae and higher plants. Two pathways of biosynthesis of the key C5-isoprene units are considered: 1) from acetate via mevalonate (C6) followed by its enzymatic conversions to isopentenyl diphosphate (C5); 2) from glucose via formation of glyceraldehyde-3-phosphate (C3) and pyruvate and their condensation via intermediary products to isopentenyl diphosphate (C5). Subsequent biosynthesis of carotenoids from isopentenyl diphosphate (C5) and dimethylallyl diphosphate (C5) involves a common route including their conversion into geranyl diphosphate (C10), farnesyl diphosphate (C15), geranylgeranyl diphosphate (C20), and synthesis of phytoene (C40). All stages of phytoene desaturation accompanied by formation of acyclic compounds such as zeta-carotene, neurosporene, and lycopene and their cyclization to alpha-, beta-, and epsilon-carotenes are considered in detail. Formation of xanthophylls in chloroplasts and chromoplasts involves sequential oxidations yielding hydroxy, epoxy, and oxo groups. Genetic control of biosynthesis of carotenoids is considered.     

Carotenoid biotechnology in plants for nutritionally improved foods

Carotenoids participate in light harvesting and are essential for photoprotection in photosynthetic plant tissues. They also furnish non-photosynthetic flowers and fruits with yellow to red colors to attract animals for pollination and dispersal of seeds. Although animals can not synthesize carotenoids de novo , carotenoid-derived products such as retinoids (including vitamin A) are required as visual pigments and signaling molecules. Dietary carotenoids also provide health benefits based on their antioxidant properties. The main pathway for carotenoid biosynthesis in plants and microorganisms has been virtually elucidated in recent years, and some of the identified biosynthetic genes have been successfully used in metabolic engineering approaches to overproduce carotenoids of interest in plants. Alternative approaches that enhance the metabolic flux to carotenoids by upregulating the production of their isoprenoid precursors or interfere with light-mediated regulation of carotenogenesis have been recently shown to result in increased carotenoid levels. Despite spectacular achievements in the metabolic engineering of plant carotenogenesis, much work is still ahead to better understand the regulation of carotenoid biosynthesis and accumulation in plant cells. New genetic and genomic approaches are now in progress to identify regulatory factors that might significantly contribute to improve the nutritional value of plant-derived foods by increasing their carotenoid levels.     

Physiology and adaptive significance of secondary carotenogenesis in green microalgae

Some species of unicellular green algae accumulate under unfavorable conditions high amounts of secondary carotenoids (SCar), which are not involved in photosynthesis and localized outside thylakoids. Such algae are cultivated on an industrial scale for production of carotenoids with valuable pharmacological properties. This review summarizes recent experimental data on the importance of secondary carotenogenesis for stress tolerance of unicellular green algae (Chlorophyta). The peculiarities of SCar biosynthesis, its induction and regulation under the action of various stressors as well as their localization in the cell and physiological effects of the accumulation of high amounts of these pigments are considered. Particular attention is given to the relationships between SCar biosynthesis and stress-induced biosynthesis of neutral lipids as well as the role of ROS as universal inducers and regulators of SCar biosynthesis. Various aspects of SCar protective function in the micfroalgal cells are discussed.     

Metabolic engineering for the microbial production of carotenoids and related products with a focus on the rare C50 carotenoids

Carotenoids, a subfamily of terpenoids, are yellow- to red-colored pigments synthesized by plants, fungi, algae, and bacteria. They are ubiquitous in nature and take over crucial roles in many biological processes as for example photosynthesis, vision, and the quenching of free radicals and singlet oxygen. Due to their color and their potential beneficial effects on human health, carotenoids receive increasing attention. Carotenoids can be classified due to the length of their carbon backbone. Most carotenoids have a C40 backbone, but also C30 and C50 carotenoids are known. All carotenoids are derived from isopentenyl pyrophosphate (IPP) as a common precursor. Pathways leading to IPP as well as metabolic engineering of IPP synthesis and C40 carotenoid production have been reviewed expertly elsewhere. Since C50 carotenoids are synthesized from the C40 carotenoid lycopene, we will summarize common strategies for optimizing lycopene production and we will focus our review on the characteristics, biosynthesis, glycosylation, and overproduction of C50 carotenoids.     

Flesh Color Diversity of Sweet Potato: An Overview of the Composition,Functions, Biosynthesis,and Gene Regulation of the Major Pigments

Sweet potato is a multifunctional root crop and a source of food with many essential nutrients and bioactive compounds. Variations in the flesh color of the diverse sweet potato varieties are attributed to the different phytochemicals and natural pigments they produce. Among them, carotenoids and anthocyanins are the main pigments known for their antioxidant properties which provide a host of health benefits, hence, regarded as a major component of the human diet. In this review, we provide an overview of the major pigments in sweet potato with much emphasis on their biosynthesis, functions, and regulatory control. Moreover, current findings on the molecular mechanisms underlying the biosynthesis and accumulation of carotenoids and anthocyanins in sweet potato are discussed. Insights into the composition, biosynthesis, and regulatory control of these major pigments will further advance the biofortification of sweet potato and provide a reference for breeding carotenoid- and anthocyanin-rich varieties.     

Endosymbiotic bacteria as a source of carotenoids in whiteflies

Although carotenoids serve important biological functions, animals are generally unable to synthesize these pigments and instead obtain them from food. However, many animals, such as sap-feeding insects, may have limited access to carotenoids in their diet, and it was recently shown that aphids have acquired the ability to produce carotenoids by lateral transfer of fungal genes. Whiteflies also contain carotenoids but show no evidence of the fungus-derived genes found in aphids. Because many sap-feeding insects harbour intracellular bacteria, it has long been hypothesized that these endosymbionts could serve as an alternative source of carotenoid biosynthesis. We sequenced the genome of the obligate bacterial endosymbiont Portiera from the whitefly Bemisia tabaci. The genome exhibits typical signatures of obligate endosymbionts in sap-feeding insects, including extensive size reduction (358.2 kb) and enrichment for genes involved in essential amino acid biosynthesis. Unlike other sequenced insect endosymbionts, however, Portiera has bacterial homologues of the fungal carotenoid biosynthesis genes in aphids. Therefore, related lineages of sap-feeding insects appear to have convergently acquired the same functional trait by distinct evolutionary mechanisms—bacterial endosymbiosis versus fungal lateral gene transfer.     

Differences in phytochrome action on the formation of carotenoids and quinones during chloroplast development in seedlings of barley (Hordeum vulgare)

The influence of phytochrome on the light induced formation of carotenoids and quinones was investigated using etiolated seedlings of barley ( Hordeum vulgare L. cv. Villa). The biosynthesis of both the quinones and the carotenoids was enhanced by active phytochrome, but the formation of the individual carotenoids and quinones was influenced by quite different thresholds. In both younger and older plants the biosynthesis of β-carotene, lutein and violaxanthin was promoted by a low P_fr threshold. The formation of plastoquinone-9, plastohydroquinone-9, α-tocoquinone and phylloquinone was also influenced by a low P_fr threshold. The biosynthesis of zeaxanthin and neoxanthin required a much higher amount of P_fr. Only antheraxanthin-likeα-tocopherol, desmethylphylloquinone and, in older leaves, α-tocoquinone exhibited a complete reversibility of the phytochrome action in their biosynthesis. The effect of phytochrome on the biosynthesis of carotenoids and quinones was different in seedlings of different age.     

2006年春夏期间浙江南麂海域浮游植物群落结构特征

研究了2006年春夏期间(4~6月)浙江省南麂列岛海域的浮游植物群落结构,共观察到68种浮游植物,其中硅藻53种,甲藻13种,金藻和蓝藻各1种。调查期间,该海域浮游植物群落结构变化明显,原甲藻赤潮的形成和持续是最显著的特征,三角棘原甲藻和东海原甲藻分别在5月中、下旬先后引发赤潮。原甲藻赤潮的形成与水温有密切关系,其生消引起海水中营养盐结构的相应变化。     

Biosynthesis of acyclic and cyclic carotenoids in higher plants and the control by phytochrome

Radish plants ( Raphanus sativus L. cv. Saxa treib) were grown in the presence of three different herbicides interfering with the biosynthesis of cyclic carotenoids. The herbicides caused an accumulation of acyclic biosynthetic intermediates. Plants were then irradiated using four different light programs in order to gain more insight into the first steps of carotenoid biosynthesis and their control by light and phytochrome. Plants grown in the dark in the presence of SAN 6706 or aminotriazole accumulated the acyclic intermediate phytoene, and those treated with J 852, the intermediates phytoene, phytofluene and zeta-carotene. In herbicide-treated plants short time irradiation with red light enhanced the formation of phytoene, phytofluene, zeta-carotene or lycopene, consistent with an effect of phytochrome on the early steps of carotenoid biosynthesis. Biosynthesis of cyclic carotenoids was also enhanced by red light in the untreated controls. In amitrole-treated plants formation of β-carotene, but not that of xanthophylls was stimulated by red light. In many cases neither the red light-induced biosynthesis of cyclic carotenoids nor the formation of acyclic intermediates could be prevented by a subsequent irradiation with far-red light. Similar enhancement as with red light was also obtained after treatment with far-red light only. Presented data may be taken as evidence that the biosynthesis and dehydrogenation of phytoene and the cyclization of lycopene are activated by a low threshold of active phytochrome. This may be further supported by the observation that far-red light itself stimulated carotenoid biosynthesis.     

The Evolution of Oxygen As a Biosynthetic Reagent

The biosynthesis of certain cell constituents: monounsaturated fatty acids, tyrosine, and nicotinic acid, is oxygen-dependent in many higher organisms. The same compounds can be synthesized by different, oxygen-independent pathways in lower organisms. The general outlines of these pathways are described and the importance of the compounds synthesized is discussed. An examination of the distribution of these pathways among living organisms reveals that oxygen-dependent pathways replaced the "anaerobic" pathways at different branch points on the evolutionary tree. Other groups of compounds are discussed, which are not distributed as widely among living organisms, but are found in all higher organisms. These compounds have specialized functions and their biosynthesis requires molecular oxygen. The oxygen-dependent portions of the biosynthetic pathways leading to porphyrins, quinone coenzymes, carotenoids, sterols, and polyunsaturated fatty acids are summarized. The distribution and functions of these compounds are also considered and an attempt is made to place them in the framework of evolution. While sterols and polyunsaturated fatty acids are found exclusively in the higher Protista and multicellular organisms, carotenoids, porphyrins, and quinones are also found in bacteria. The possibility of oxygen-independent mechanisms for their biosynthesis is discussed.     

安徽沱湖夏季浮游植物群落结构特征与环境因子关系

为了揭示沱湖浮游植物群落结构特征及其与水环境因子的关系,于2016年7月(夏季),对沱湖流域上游至下游11个采样点浮游植物种类组成、细胞丰度、生物量等进行调查研究。结果显示,沱湖共有浮游植物96种(含变种),隶属8门48属,其中绿藻门(Chlorophyta)和硅藻门(Bacillariophyta)种类最多,绿藻门有23属39种,占总种数的40.63%,硅藻门有7属20种,占总种数的20.83%;其次为裸藻门(Euglenophyta),有5属17种,占总种数的17.71%,蓝藻门(Cyanophyta) 8属14种,占14.58%;甲藻门(Pyrrophyta) 2属2种,隐藻门(Cryptophyta) 1属2种,各占总种数的2.08%,黄藻门(Xanthophyta)与金藻门(Chrysophyta)均有1属1种,均占总种数的1.04%。绿藻和硅藻类物种在沱湖浮游植物群落结构中处于优势地位,沱湖夏季浮游植物种类组成表现为绿藻-硅藻型。沱湖夏季浮游植物细胞丰度与生物量从上游到下游呈逐渐增加趋势,细胞丰度与生物量平均值分别为4.022×106cells/L与3.046 mg/L,蓝藻门和绿藻门类群为沱湖浮游植物细胞丰度主体,硅藻门和裸藻门类群为沱湖浮游植物生物量的主体;上游浮游植物多样性指数与均匀度指数均略高于下游采样点,沱湖水质呈β中污型-无污染型,上游水质优于下游水质。浮游植物群落结构与水环境因子的典型对应分析(CCA)结果表明,电导率、透明度、水深及pH值等环境因子与沱湖夏季浮游植物群落结构有较强的相关性。     

Making extra room for carotenoids in plant cells: New opportunities for biofortification

Plant carotenoids are essential for photosynthesis and photoprotection and provide colors in the yellow to red range to non-photosynthetic organs such as petals and ripe fruits. They are also the precursors of biologically active molecules not only in plants (including hormones and retrograde signals) but also in animals (including retinoids such as vitamin A). A carotenoid-rich diet has been associated with improved health and cognitive capacity in humans, whereas the use of carotenoids as natural pigments is widespread in the agrofood and cosmetic industries. The nutritional and economic relevance of carotenoids has spurred a large number of biotechnological strategies to enrich plant tissues with carotenoids. Most of such approaches to alter carotenoid contents in plants have been focused on manipulating their biosynthesis or degradation, whereas improving carotenoid sink capacity in plant tissues has received much less attention. Our knowledge on the molecular mechanisms influencing carotenoid storage in plants has substantially grown in the last years, opening new opportunities for carotenoid biofortification. Here we will review these advances with a particular focus on those creating extra room for carotenoids in plant cells either by promoting the differentiation of carotenoid-sequestering structures within plastids or by transferring carotenoid production to the cytosol.     

植物类胡萝卜素生物合成及功能

详述了植物类胡萝卜素生物合成途径,并从突破类胡萝卜素合成途径中上游瓶颈限制、类胡萝卜素代谢各分支途径的改造、提高植物细胞对类胡萝卜素物质积累能力三个方面探讨了类胡萝卜素生物合成酶基因在植物基因工程中的研究现状,最后对植物类胡萝卜素代谢的研究前景进行了展望。     

Diversity of microbial eukaryotes in Kongsfjorden,Svalbard

Microbial eukaryote diversity was assessed in Arctic Kongsfjorden (Svalbard), by constructing SSU rDNA clone libraries. Samples were collected from different depths at the outer basin in summer (2006), plus an additional one glacial and one sediment sample. The libraries displayed diversity based on 284 full-length sequences. Four main phyla, namely, Alveolates, Stramenopiles, Cercozoans, and Metazoans were often screened in this fjord. Alveolate occupied the highest percentage of taxa in the library of surface sea water, besides the Metazoan-related clones. Moreover, dinoflagellates, diatoms, and pico-Prasinophytes were detected as prevalent phytoplankton through the analysis of libraries. Questions related to the quantity of these phytoplankton and their roles in the microbial food loop arose from an ecological viewpoint.     

A RING-finger photocarotenogenic repressor involved in asexual sporulation in Mucor circinelloides.

Mucor circinelloides responds to blue light by activating the biosynthesis of carotenoids and bending its sporangiophores towards the light source. The CrgA protein product acts as a repressor of carotene biosynthesis, as its inactivation leads to the overaccumulation of carotenoids in both the dark and the light. We show here that asexual sporulation in Mucor is also stimulated by light and that the crgA gene is involved in sporulation, given that lack of crgA function affects both carotenogenesis and the normal production of spores. A small interference RNA (siRNA) gene silencing approach was used to block the biosynthesis of carotenoids and to demonstrate that abnormal sporulation in crgA mutants is not a consequence of a defective production of carotenes. These results reveal an active role for the predicted CrgA product, a RING-finger protein, in the control of cellular light-regulated processes in Mucor.     

Carotenogenesis diversification in phylogenetic lineages of Rhodophyta

Carotenoid composition is very diverse in Rhodophyta. In this study, we investigated whether this variation is related to the phylogeny of this group. Rhodophyta consists of seven classes, and they can be divided into two groups on the basis of their morphology. The unicellular group (Cyanidiophyceae, Porphyridiophyceae, Rhodellophyceae, and Stylonematophyceae) contained only β‐carotene and zeaxanthin, “ZEA‐type carotenoids.” In contrast, within the macrophytic group (Bangiophyceae, Compsopogonophyceae, and Florideophyceae), Compsopogonophyceae contained antheraxanthin in addition to ZEA‐type carotenoids, “ANT‐type carotenoids,” whereas Bangiophyceae contained α‐carotene and lutein along with ZEA‐type carotenoids, “LUT‐type carotenoids.” Florideophyceae is divided into five subclasses. Ahnfeltiophycidae, Hildenbrandiophycidae, and Nemaliophycidae contained LUT‐type carotenoids. In Corallinophycidae, Hapalidiales and Lithophylloideae in Corallinales contained LUT‐type carotenoids, whereas Corallinoideae in Corallinales contained ANT‐type carotenoids. In Rhodymeniophycidae, most orders contained LUT‐type carotenoids; however, only Gracilariales contained ANT‐type carotenoids. There is a clear relationship between carotenoid composition and phylogenetics in Rhodophyta. Furthermore, we searched open genome databases of several red algae for references to the synthetic enzymes of the carotenoid types detected in this study. β‐Carotene and zeaxanthin might be synthesized from lycopene, as in land plants. Antheraxanthin might require zeaxanthin epoxydase, whereas α‐carotene and lutein might require two additional enzymes, as in land plants. Furthermore, Glaucophyta contained ZEA‐type carotenoids, and Cryptophyta contained β‐carotene, α‐carotene, and alloxanthin, whose acetylenic group might be synthesized from zeaxanthin by an unknown enzyme. Therefore, we conclude that the presence or absence of the four enzymes is related to diversification of carotenoid composition in these three phyla.     

异戊二烯类植物激素赤霉素和脱落酸的代谢调控

赤霉素和脱落酸在植物生理过程中具有重要的调控作用,其生物合成途径迄今已基本阐明。赤霉素与类胡萝卜素的生物合成途径具有共同前体牻牛儿基牻牛儿基二磷酸,而脱落酸则直接来自于类胡萝卜素。参与这两种植物激素和类胡萝卜素代谢过程的大多数酶基因已经从不同植物中获得克隆;各种调控方式也随着分子生物学的研究工作而得到鉴定。本文就近年来对赤霉素和脱落酸等代谢调控机制及其与植物类胡萝卜素代谢之间关系的研究工作做简要回顾。     

The role of phytoplankton in the diet of the bladderwort Utricularia australis R.Br. (Lentibulariaceae)

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