基于18S rRNA基因序列分析唇口目苔藓动物主要类群的系统发生关系

对隶属于3亚目、5次目、20科、23属共25个种类的唇口目(裸唇纲)苔藓虫18S rRNA基因部分序列进行了序列测定.结合从GenBank中获得的该类群其它7个种类的18S rRNA基因同源序列,以序列分析软件对其序列组成和变异进行了比较分析;同时,以羽苔虫(被唇纲)和管孔苔虫(窄唇纲)为外类群,以邻接法和最大简约法重建了它们的系统发生树,分析了该目主要类群系统发生关系.序列分析结果显示:经比对后序列长度为884 bp,其中保守位点241个,可变位点643个,简约信息位点357个;A,T,C和G 4碱基平均含量分别为23.8%、22.8%、24.4%和28.9%.分子系统树表明:本研究所有有囊类构成1个单系群,其中檐胞次目的几种苔虫位于皮壳次目内部;无囊类形成1个多系群,其中的亚目级(新唇口亚目)和次目级分类阶元(枝室次目、假软壁次目和隐壁次目)也都为多系发生,这些结果与前人的分子系统学研究结果大体一致,而与传统的形态分类体系间存在明显的冲突.     

基于16S rRNA基因序列探讨中国粉蛉科的系统发育关系

本研究测定了粉蛉科Coniopterygidae9属15种昆虫16SrRNA基因部分序列,对序列的碱基组成、转换/颠换比率、遗传距离、变异位点等进行分析。基于16SrRNA基因序列数据,分别采用邻接法（NJ）、最大简约法（MP）和最大相似法（ML）建立粉蛉科分子系统发育关系。研究结果表明：粉蛉亚科的粉蛉属Coniopteryx与重粉蛉属Semilalis是姊妹群,（虫齿）粉蛉属Conwentzia较前二者原始;囊粉蛉亚科的卷粉蛉属Helicoconis和隐粉蛉属Cryptoscenea的亲缘关系较近。曲粉蛉属Coniocompsa和异粉蛉属Heteroconis聚类在一起,但自展值较低。瑕粉蛉属Spiloconis和囊粉蛉属Aleuropteryx的位置在囊粉蛉亚科中不够稳定。     

介形类系统分类与起源演化的形态、分子和化石证据

介形类(Ostracoda)因其丰富的化石记录和广布的海陆现生代表类群,而被认为是进化生物学中研究生物多样性产生机制和演变历程的颇具潜力的重要模式生物。介形类在甲壳亚门中的谱系发生位置、起源及其内部各类群间的系统关系还存在诸多争议。基于其体制构造的形态学特征,介形类被归入甲壳亚门下的颚足纲(Maxillopoda),但来自18S rDNA序列数据分析却显示Maxillopoda不是单系群。基于化石记录和壳体形态特征,高肌虫(Bradoriida)长期以来被认为是介形类的一个祖先类群,但保存有软躯体的早寒武世化石的研究表明,Bradoriida不是介形类甚至可能也不属于甲壳类。不同的研究者所强调的壳体和肢体形态特征各异,导致介形类最大的现生类群速足目(Podocopida)的四个超科之间的关系也存在诸多推测。壳体和肢体特征在系统演化意义上的不兼容,需要分子生物学等证据的介入。分子、形态和化石证据的积累及各种信息整合是系统演化研究的必然趋势。     

PHYLOGENETIC AFFINITIES OF CRYSONEPHELE PALUSTRIS (CHRYSOPHYCEAE) BASED ON INFERRED NUCLEAR SMALL-SUBUNIT RIBOSOMAL RNA SEQUENCE

The nuclear small-subunit ribosomal DNA sequence was determined from the colonial chromophytic alga Chrysonephele palustris Pipes, Tyler, et Leedale using polymerase chain reaction methods. The inferred ribosomal RNA sequence was included in a multiple alignment, containing heterokont chromophytes, and subjected to molecular systematic analyses in order to determine the phylogenetic relationships of this alga . Chrysonephele palustris grouped within the Chrysophyceae and not in an intermediate position between the Chrysophyceae and Eustigmatophyceae, suggesting that Chrysonephele is not a phylogenetic link between these algal classes .     

A PHYLOGENETIC ANALYSIS OF THE SYNUROPHYCEAE USING MOLECULAR DATA AND SCALE CASE MORPHOLOGY

The phylogeny of the Synurophyceae was investigated by parsimony analysis of scale case characters and small-sub unit (18S) ribosomal RNA (rRNA) sequence data. Analysis of 1 eustigmatophycean (outgroup), 3 chrysophycean, and 10 synurophycean 18S rRNA sequences corroborated the inference from ultrastructural information that the Synurophyceae is a monophyletic assemblage . Tessellaria vol-vocina, which had been tentatively proposed as a member of the Synurophyceae, was confirmed as the earliest lineage within the Synurophyceae by both the molecular analysis and an evaluation of published ultrastructural data. A second set of analyses investigated the relationships among Tessellaria volvocina, 6 Synura species, and 10 Mallomonas species/varieties, with particular reference to the validity of current classifications of the Synurophyceae and the characters upon which they are based. The molecular and scale case phylogenies were not totally resolved but were largely congruent. The data sets were combined to produce another phylogeny, which showed greater resolution. The combined phylogeny weakly supported our representatives of Synura and Mallomonas as monophyletic groupings and also upheld several of the sections within these genera that are recognized by current classifications. However, some changes to the classification and delineation of these genera are recommended and predicted. Both our 18S rRNA sequence and scale case data sets were more appropriate for examining the branching order among the more closely related text rather than resolving the deeper branching points of the synurophycean phylogeny .     

PHYLOGENETIC ANALYSIS OF YAMAGISHLELLA AND PLATYDORINA (VOLVOCACEAE,CHLOROPHYTA) BASED ON rbcL GENE SEQUENCES1

Yamagishiella, based on Pandorina unicocca Rayburn et Starr, is distinguished from Eudorina by its isogamous sexual reproduction, whereas Platydorina exhibits anisogamous sexual reproduction. In the present study, we sequenced the large subunit of ribulose-1,5-bisphosphate carboxylase/oxygenase (rbcL) genes from five Japanese and North American strains of Y. unicocca (Rayburn et Starr) Nozaki, two Platydorina caudata Kofoid strains, and two strains of Eudorina unicocca G. M. Smith, as well as eight related colonial and unicellular species. Phylogenetic trees were constructed based on these sequence data and on previously published rbcL gene sequences from 23 volvocalean species in order to deduce phylogenetic relationships within the colonial Volvocales, with particular regard to the phylogenetic positions and status of the genera Yamagishiella and Platydorina. Two robust monophyletic groups of the anisogamous/oogamous volvocacean species were resolved in the maximum-parsimony tree as well as in the neighbor-joining distance tree. One of the two groups comprises three species of Volvox section Volvox, whereas the other is composed of other sections of Volvox as well as of all the species of Eudorina and Pleodorina. Platydorina, however, was positioned outside these two monopliyletic groups. Therefore, derivation of the Platydorina lineage may be earlier than that of such anisogamous/oogamous groups, or orgin of “anisogamy with sperm packets” in Platydorina may be independent of sperm packet evolution in Eudorina, Pleodorina, and Volvox. It was also resolved with high bootstrap values that all of the Y. unicocca strains form a monophyletic group positioned outside the large monophyletic group including Eudorina and Pleodorina. These reject the possibility of the reverse evolution of isogamy from anisogamy to give rise to Yamagishiella within the lineage of Eudorina.     

A PHYLOGENETIC STUDY OF THE CORALLINALES (RHODOPHYTA) BASED ON NUCLEAR SMALL-SUBUNIT rRNA GENE SEQUENCES

Conflicting classifications for the Corallinales were tested by analyzing partial sequences for the nuclear small-subunit ribosomal RNA (SSU) gene of 35 species of coralline algae. Parsimony and likelihood analyses of these data yielded congruent hypotheses that are inconsistent with classifications for the group that include as many as eight subfamilies. Four major clades are resolved within the order, including the early-diverging Sporolithaceae as well as the Melobesioideae and Corallinoideae. The fourth clade, which is supported robustly, includes both nongeniculate and geniculate species classified in the subfamilies Mastophoroideae, Metagoniolithoideae, Lithophylloideae, and Amphiroideae. Molecular and morphological data support the proposal that the latter two subfamilies are sister taxa. Although relationships among some genera are not resolved clearly, the order of branching of taxa among and within the four principal lineages is concordant with paleontological evidence for the group. Relationships inferred among genera within each of the clades is discussed. Seven morphological characters delimiting higher taxonomic groups within the order were combined with the sequence data, analyzed, and optimized onto the resulting tree(s). Except for the presence or absence of genicula, all other characters were found to be phylogenetically informative. Genicula are nonhomologous structures that evolved independently in the Amphiroideae, Corallinoideae, and Metagoniolithoideae. The phenetic practice of separating coralline algae into two categories solely on the basis of the presence or absence of genicula does not accurately reflect the evolutionary history of the group.     

PHYLOGENETIC RELATIONSHIPS OF THE FRESHWATER ALGA BOLDIA ERYTHROSIPHON (COMPSOPOGONALES, RHODOPHYTA) BASED ON 18S rRNA GENE SEQUENCES

The nuclear small-subunit ribosomal DNA sequence from the freshwater red alga Boldia erythrosiphon Herndon emend Howard et Parker was determined. Phylogenetic analysis confirms the positioning of this species within the bangiophycidean order of the Compsopogonales. The results strongly suggest that occupation of freshwater habitats by marine red algae has happened at least twice during red algal evolution.     

几种蚊虫线粒体DNA-16SrRNA序列及其相互关系的研究

测定我国尖音库蚊复合组4亚种（尖音库蚊、淡色库蚊、致倦库蚊和骚扰库蚊）、三带喙库蚊、白纹伊蚊和中华按蚊的线粒体DNA 16S rRNA基因（mtDNA-16S rRNA)序列,发现我国尖音库蚊复合组4亚种mtDNA-16S rRNA序列基本一致,长度为555bp(554bp),GC含量为25．41％。该序列与其他3种蚊虫在种间存在差异,与三带喙库蚊的种间差异为0．54％;与白纹伊蚊的种间差异为5．77％;与中华按蚊的种间差异为9．62％。分子系统关系分析表明该序列与传统分类系统的同属或同亚科种类近似性相一致。     

ANALYSIS OF CLONAL CULTURES OF THE BROWN TIDE ALGAE AUREOCOCCUS AND AUREOUMBRA (PELAGOPHYCEAE) USING 18S rRNA, rbcL, AND RUBISCO SPACER SEQUENCES

Variability among clonal cultures of the brown tide algae Aureococcus anophagefferens Hargraves et Sieburth and Aureoumbra lagunensis Stockwell, DeYoe, Hargraves et Johnson was examined by DNA sequence comparisons. Nuclear-encoded 18S rRNA and plastid-encoded rbc L gene sequences were determined for six Aureococcus strains. RUBISCO spacer sequences were determined for 14 strains of Aureococcus. No differences among Aureococcus strains were found in the DNA regions examined. The rbc L and RUBISCO spacer sequences for three Aureoumbra strains were identical but differed from those of Aureococcus. These data indicate that blooms of these species are comprised of cells that are very similar and also imply that Aureococcus and Aureoumbra do not contain varieties or cryptic species. Separate and combined phylogenetic analyses of the 18S rRNA and rbc L gene sequences were performed. Results confirm that the brown-tide-causing algae of Long Island Sound, New York ( Aureococcus ), and Laguna Madre, Texas ( Aureoumbra ), are best classified in separate genera within the Pelagophyceae. Phylogenetic trees place Aureococcus and Aureoumbra within the Pelagomonadales and Sarcinochrysidales, respectively.     

基于16S rRNA基因序列对织纹螺属的分子系统学分析

研究利用线粒体16S rRNA基因序列片段,对织纹螺属6亚属11种动物进行分子系统学分析。结果显示,种内遗传距离(0～0.007)与种间遗传距离(0.019～0.088)无重叠,这表明线粒体16S rRNA基因能较好地反映织纹螺种间的亲缘关系; 而亚属内遗传距离(0.018～0.031)与亚属间遗传距离(0.028～0.083)存在重叠,表明线粒体16S rRNA基因不能对一些贝壳形态相似的亚属进行区分。同时,确定了疑难种灰白织纹螺Nassarius(Zeuxis)canaliculatum的分类地位,并与相似种西格织纹螺N.(Z.)siquijorensis进行了对比; 确认了秀丽织纹螺Nassarius(Hima)festivus应属于Hima亚属;建议保留单型亚属Varicinassa的有效性。     

基于18S rRNA基因序列的直翅目主要类群系统发育关系研究

基于78种直翅目昆虫的18S rRNA基因全序列构建了直翅目各主要类群间的系统发育关系。本研究的结果支持直翅目的单系性,但不支持蝗亚目和螽亚目各自的单系性;直翅目下除蜢总科和蝗总科外各总科的划分多数与Otte系统相一致;蜢总科的单系性得不到支持;蝗总科的剑角蝗科、斑腿蝗科、斑翅蝗科、网翅蝗科和槌角蝗科5科均不是单系群,各物种间的遗传距离差异不大,应合并为一科,即蝗科;本研究支持将Otte系统中蚱总科和螽蟖总科下各亚科级阶元提升为科级阶元;18S rRNA基因全序列可以作为划分科级阶元的工具,当位于同一分支上互成姐妹群的类群间的遗传距离超过1%时,这几个类群属于不同的科;但由于其在进化上的保守性,18S rRNA基因只能用于纲目等高级阶元间关系的研究,而由其获得的总科以下阶元间的关系并不可靠。     

基于发生学与形态学资料对盾纤目纤毛虫(原生动物,纤毛门)的系统修订

根据细胞分裂中口器发生的类型、形态结构特征及近祖-衍化性状的推断,利用简约性原理及聚类分析方法对盾纤目纤毛虫的目下系统关系做了探讨和修订.主要结论包括:1)确认该类群为一目级的单源发生系,下含3个亚目:嗜污亚目、帆口亚目、触毛亚目;2)嗜污亚目经廓清认定了52个有效属,分别隶属重新调整后的13个科;3)帆口亚目经厘定共有20个有效属,调整入7个科中;4)触毛亚目3个科,包含13个地位明确的属.文中对前人所建系统中不尽合理、存疑以及混乱之处做了讨论和建议性修正.     

苔藓动物18S rRNA基因的分子系统发生初探

本文对我国沿海较为常见的8种唇口目苔藓动物的18SrRNA基因进行了PCR扩增和序列测定。结合已知的其它苔藓动物（包括内肛动物和外肛动物）以及腕足动物和帚虫的相应序列,运用分子系统学方法,研究苔藓动物门的系统发生关系,结果表明,外肛动物和内肛动物构成苔藓动物分子系统树中的二大平行支;本文测定的大室膜孔苔虫与Giribet等测定的膜孔苔虫在系统树中的位置间隔较远。结果也支持外肛动物包含被唇纲和裸唇纲两大类群的形态划分,而关于裸唇纲特别是唇口目内部的系统发生关系。分子数据的分析结果和形态分类之间的分歧有待于进一步研究。     

MOLECULAR PHYLOGENETIC ANALYSIS OF THE HETEROTROPHIC CHRYSOPHYTE GENUS PARAPHYSOMONAS (CHRYSOPHYCEAE), AND THE DESIGN OF rRNA-TARGETED OLIGONUCLEOTIDE PROBES FOR TWO SPECIES

Nanoflagellate protists are algae and protozoa (2– 20 μm in size) that play important ecological roles in freshwater and marine microbial communities as primary producers and as consumers of prokaryotic and eukaryotic prey. There is little biogeographical information for most of these minute protists despite their significant role in aquatic food webs. In addition, the evolutionary relationships among some of these species and their affinities to other protistan taxa are unclear. These circumstances are largely a consequence of the fact that small protists possess few readily apparent morphological features on which to base taxonomic and phylogenetic schemes and with which to identify them in natural assemblages. As an alternative approach for addressing these issues, we sequenced the small-subunit ribosomal RNA genes of four species of the colorless chrysophyte genus Paraphysomonas. A phylogenetic analysis based on that sequence information was performed, and oligonucleotide probes for two commonly occurring species of Paraphysomonas were designed and tested. Phylogenetic analyses of these four species confirmed the affinity of the genus Paraphysomonas with other chrysophyte species. High sequence similarity among three of the species (P. imperforata Lucas, P. bandaiensis Takahashi, and P. foraminifera Lucas) supported a previous phylogenetic grouping of these species based on the morphology of the scales produced by these species. In particular, sequence similarity between P. imperforata and P. foraminifera indicated that this speciation was a recent evolutionary event. However, a fourth species (P. vestita (Stokes) de Saedeleer) possessing similar scale morphology to P. bandaiensis, P. imperforata, and P. foraminifera showed considerable sequence dissimilarity in comparison to these latter three species. Oligonucleotide probes were successfully designed for the species P. imperforata and P. bandaiensis and applied together with a recently developed quantitative in situ hybridization procedure. The development of species-specific oligonucleotide probes for these nanoflagellate species and their application for counting nanoflagellates in natural water samples provide tools for studying these ecologically important species.     

PHYLOGENY OF CARTERIA (CHLOROPHYCEAE) INFERRED FROM MOLECULAR AND ORGANISMAL DATA1

Comparative ultrastructural data have shown that at least two distinct groups exist within Carteria. Similarly, interpretations of variation in gross morphological features have led to the discovery of morphologically distinct groups within the genus. Partial sequences from the nuclear-encoded small- and large-subunit ribosomal RNA molecules of selected Carteria taxa were studied as a means of 1) testing hypotheses that distinct groups of species exist within the genus and 2) assessing monophyly of the genus. Parsimony analysis of the sequence data suggests that three Carteria species, C. lunzensis, C. crucifera, and C. olivieri, form a monophyletic group that is the basal sister group to all other ingroup flagellate taxa (including species of Chlamydomonas, Haematococcus, Stephanosphaera, Volvox, and Eudorina). Two other Carteria taxa, C. radiosa and Carteria sp. (UTEX isolate LB 762), form a clade that is the sister group to a clade that includes Haematococcus spp., Chlamydomonas spp., and Stephanosphaera. Thus, the sequence data support the interpretations of ultrastructural evidence that described two distinct Carteria lineages. Moreover, the sequence data suggest that these two Carteria groups do not form a monophyletic assemblage. Parsimony analysis of a suite of organismal (morphological, ultra-structural, life history, and biochemical) character data also suggest two distinct lineages among the five Carteria taxa; however, the organismal data are ambiguous regarding monophyly of these Carteria taxa. When the two independent data sets are pooled, monophyly of Carteria is not supported; therefore, the weight of available evidence, both molecular and organismal, fails to support the concept of Carteria as a natural genus.     

UPDATING THE GENUS DICTYOSPHAERIUM AND DESCRIPTION OF MUCIDOSPHAERIUM GEN. NOV. (TREBOUXIOPHYCEAE) BASED ON MORPHOLOGICAL AND MOLECULAR DATA1

Recent molecular analyses of Dictyosphaerium strains revealed a polyphyletic origin of this morphotype within the Chlorellaceae. The type species Dictyosphaerium ehrenbergianum Nägeli formed an independent lineage within the Parachlorella clade, assigning the genus to this clade. Our study focused on three different Dictyosphaerium species to resolve the phylogenetic position of remaining species. We used combined analyses of morphology; molecular data based on SSU and internally transcribed spacer region (ITS) rRNA sequences; and the comparison of the secondary structure of the SSU, ITS‐1, and ITS‐2 for species and generic delineation. The phylogenetic analyses revealed two lineages without generic assignment and two distinct clades of Dictyosphaerium‐like strains within the Parachlorella clade. One clade comprises the lineages with the epitype strain of D. ehrenbergianum Nägeli and two additional lineages that are described as new species (Dictyosphaerium libertatis sp. nov. and Dictyosphaerium lacustre sp. nov.). An emendation of the genus Dictyosphaerium is proposed. The second clade comprises the species Dictyosphaerium sphagnale Hindák and Dictyosphaerium pulchellum H. C. Wood. On the basis of phylogenetic analyses, complementary base changes, and morphology, we describe Mucidosphaerium gen. nov with the four species Mucidosphaerium sphagnale comb. nov., Mucidosphaerium pulchellum comb. nov., Mucidosphaerium palustre sp. nov., and Mucidosphaerium planctonicum sp. nov.     

CHARACTERIZATION AND PHYLOGENETIC POSITION OF THE ENIGMATIC GOLDEN ALGA PHAEOTHAMNION CONFERVICOLA: ULTRASTRUCTURE, PIGMENT COMPOSITION AND PARTIAL SSU rDNA SEQUENCE

The morphology, ultrastructure, photosynthetic pigments, and nuclear-encoded small subunit ribosomal DNA (SSU rDNA) were examined for Phaeothamnion confervicola Lagerheim strain SAG119.79. The morphology of the vegetative filaments, as viewed under light microscopy, was indistinguishable from the isotype. Light microscopy, including epifluorescence microscopy, also revealed the presence of one to three chloroplasts in both vegetative cells and zoospores. Vegetative filaments occasionally transformed to a palmelloid stage in old cultures. An eyespot was not visible in zoospores when examined with light microscopy, but small droplets, similar to eyespot droplets, were apparent beneath the shorter flagellum when cells were viewed with electron microscopy. Zoospores had two flagella that were laterally inserted in the cell approximately one-third of the cell length from the apex. The longer flagellum was directed anteriorly and the shorter flagellum was directed posteriorly. Electron microscopy revealed the presence of tubular tripartite flagellar hairs on the longer flagellum, but no lateral filaments were found on the tripartite hairs. The general organization of the flagellar root system was similar to that of zoospores belonging to the Xanthophyceae and Phaeophyceae. However, the transitional region of the flagella contained a transitional helix with four to six gyres. Microtubular root R₁ consisted of six microtubules at its proximal end and one microtubule at its distal end. Roots R₂ and R₄ consisted of one microtubule each and root R₃ consisted of two microtubules. No rhizoplast was found. Thin-layer chromatography revealed the presence of fucoxanthin, diadinoxanthin, neoxanthin, and heteroxanthin as well as chlorophylls a, c₁ and c₂. High-performance liquid chromatography revealed the presence of fucoxanthin, diadinoxanthin, diatoxanthin, heteroxanthin, and β,β-carotene as well as chlorophylls a and c. The complete sequence of the SSU rDNA could not be obtained, but a partial sequence (1201 bases) was determined. Parsimony and neighbor-joining distance analyses of SSU rDNA from Phaeothamnion and 36 other chromophyte algae (with two Öomycete fungi as the outgroup) indicated that Phaeothamnion was a weakly supported (bootstrap = <50%, 52%) sister taxon to the Xanthophyceae representatives and that this combined clade was in turn a weakly supported (bootstrap = <50%, 67%) sister to the Phaeophyceae. Based upon ultrastructural observations, pigment analysis, and SSU rDNA phylogenetic analysis, Phaeothamnion is not a member of the Chrysophyceae and should be classified as incertae sedis with affinities to the Xanthophyceae and Phaeophyceae.     

MOLECULAR EVOLUTION OF EUGLENOZOAN PARAXONEMAL ROD GENES par1 AND par2 COINCIDES WITH PHYLOGENETIC RECONSTRUCTION BASED ON SMALL SUBUNIT rDNA DATA1

Emergent flagella of Euglenozoa consist of two prominent structural elements: the axoneme built by microtubules with motor proteins to enable the movement of the flagellum and a highly organized protein structure of unknown function, called the paraxonemal rod (PAR), which consists of two major proteins paralleling the axoneme of euglenid and kinetoplastid emergent flagella. These flagellar structures are considered apomorphic characters of Euglenozoa. We examined the evolution of the genes par1 and par2 encoding the two major proteins, where we could show that these proteins are encoded by two very similar genes found in kinetoplastids and euglenids. The branching pattern indicated a gene duplication before the diversification into euglenids and kinetoplastids. In the clades of the genes, subtrees of euglenid and kinetoplastid monophyla arose. Both genes showed strong genetic diversity with biased GC content at taxon rather than at gene level. We also examined phylogenies inferred from PAR genes that are well in agreement with established small subunit rDNA analyses. Both showed further separation of the euglenid subtree into primary osmotrophs and a phototrophic clade, including secondarily derived osmotrophs.     

CHARACTERIZATION OF THE LIGHT-HARVESTING COMPLEX OF GIRAUDYOPSIS STELLIFER (CHRYSOPHYCEAE) AND EFFECTS OF LIGHT STRESS1

The light-harvesting complex (LHC) of Giraudyopsis stellifer Dangeard was isolated on a sucrose density gradient after digitonin treatment, and the pigment composition was analyzed by reverse-phase high-pressure liquid chromatography. The LHC is a chlorophyll (Chl) a/c/ fucoxanthin/violaxanthin complex, depleted of β-carotene, comparable to the LHC of Fucophyceae. The excitation transfer from Chl c and fucoxanthin to Chl a is efficient in whole cells. Immunological reactions indicate a close relationship between Chrysophyceae and Fucophyceae. The immunocytochemical labeling confirms the lack of segregation of the LHC in the appressed membranes of the three associated thylakoids and its localization in the intrapyrenoid thylakoid. The violaxanthin-antheraxanthin-zeaxanthin cycle is operative in the cells and efficiently protects photosystem II reaction centers against photoinhibition.