Schmitt R

Differention of germinal and somatic cells in Volvox carteri
CURRENT OPINION IN MICROBIOLOGY 6 (6): 608-613 DEC 2003


Abstract:
Volvox carteri is a spherical alga with a complete division of labor between around 2000 biflagellate somatic cells and 16 asexual reproductive cells (gonidia). It provides an attractive system for studying how a molecular genetic program for cell-autonomous differentiation is encoded within the genome. Three types of genes have been identified as key players in germ-soma differentiation: a set of gls genes that act in the embryo to shift cell-division planes, resulting in asymmetric divisions that set apart the large-small sister-cell pairs; a set of lag genes that act in the large gonidial initials to prevent somatic differentiation; and the regA gene, which acts in the small somatic initials to prevent reproductive development. Somatic-cell-specific expression of regA is controlled by intronic enhancer and silencer elements.

 

Nozaki H

Origin and evolution of the genera Pleodorina and Volvox (Volvocales)
BIOLOGIA 58 (4): 425-431 JUL 2003

 

Abstract:
The previous molecular phylogenetic study using 6021 base pairs from five chloroplast genes suggested that two species of Pleodorina (P. californica, P. japonica) might have evolved from a Volvox-like alga by the decrease in colony cell number and size. However, number of species of the genus Volvox was very limited especially in the section Merrillosphaera. In the present study, 6021 base pairs of the concatenated five chloroplast genes from 10 strains representing seven taxa of the genus Volvox were added to the previous data matrix. The sequence data resolved two anisogamous/oogamous clades within a large monophyletic group comprising five advanced genera of the Volvocaceae (Yamagishiella, Platydorina, Eudorina, Pleodorina and Volvox), one containing Volvox sect. Volvox and the anisogamous genus Platydorina (32-celled flattened colony), and the other (Eudorina group) composed of three other sections of Volvox, Pleodorina and Eudorina. The isogamous genus Yamagishiella (32-celled colony) was positioned basally to the Eudorina group. Therefore, evolution of anisogamy with sperm packets from isogamy might have occurred twice within the Volvocaceae. Based on the present molecular phylogenetic analysis, species of Volvox and Pleodorina within the Eudorina group represented three and two, respectively, separate lineages. One the three Volvox lineages [composed of V (sect. Merrillosphaera) carteri, V (sect. Merrillosphaera) obversus, V. (sect. Merrillosphaera) tertius, V. (sect. Merrillosphaera) africanus and V (sect. Copelandosphaera) dissipatrix] was sister to the monophyletic group consisting of one of the two Pleodorina lineages (P. californica and P. japonica) and V (sect. Janetosphaera) aureus. In addition, species of Eudorina were basal to the two lineages of Pleodorina and three Volvox lineages within the Eudorina group, representing the ancestral situation of Pleodorina/Volvox (excluding sect. Volvox). Thus, reverse evolution from a Volvox-like alga to Pleodorina suggested previously appears unlikely.

 

Cheng Q, Fowler R, Tam LW, et al.

The role of GlsA in the evolution of asymmetric cell division in the green alga Volvox caiteri
DEV GENES EVOL 213 (7): 328-335 JUL 2003

 

Abstract:
Volvox carteri, a green alga in the order Volvocales, contains two completely differentiated cell types, small motile somatic cells and large reproductive cells called gonidia, that are set apart from each other during embryogenesis by a series of visibly asymmetric cell divisions. Mutational analysis has revealed a class of genes (gonidialess, gls) that are required specifically for asymmetric divisions in V. carteri, but that are dispensable for symmetric divisions. Previously we cloned one of these genes, glsA, and showed that it encodes a chaperone-like protein (G1sA) that has close orthologs in a diverse set of eukaryotes, ranging from fungi to vertebrates and higher plants. In the present study we set out to explore the role of glsA in the evolution of asymmetric division in the volvocine algae by cloning and characterizing a glsA ortholog from one of the simplest members of the group, Chlamydomonas reinhardtii, which does not undergo asymmetric divisions. This ortholog (which we have named gar1, for glsA related) is predicted to encode a protein that is 70% identical to G1sA overall, and that is most closely related to G1sA in the same domains that are most highly conserved between G1sA and its other known orthologs. We report that a gar1 transgene fully complements the glsA mutation in V. carteri, a result that suggests that asymmetric division probably arose through the modification of a gene whose product interacts with G1sA, but not through a modification of glsA itself.

 

Grewing A, Krings M, Galtier J, et al.

The oldest fossil endophytic alga and its unusual habitat
SYMBIOSIS 34 (3): 215-230 2003

Abstract:
Lycophyte megaspores from the Lower Carboniferous of France sometimes contain a colonial (volvocacean) alga as an endophyte. This peculiar plant-plant association was briefly described more than 100 years ago and the name Lageniastrum macrosporae introduced for the alga, but the biological significance of the discovery was never fully appreciated. Here we present a reappraisal of the original material, which to date provides the oldest unequivocal fossil evidence for endophytic algae and the only example of an alga residing in the interior of spores of vascular cryptogams.

Nedelcu AM, Michod RE

Sex as a response to oxidative stress: the effect of antioxidants on sexual induction in a facultatively sexual lineage
P ROY SOC LOND B BIO 270: S136-S139 Suppl. 2
NOV 7 2003 (for pdf click here)

 

Abstract:
The evolution of sex is one of the long-standing unsolved problems in biology. Although in many lineages sex is an obligatory part of the life cycle and is associated with reproduction, in prokaryotes and many lower eukaryotes, sex is facultative, occurs in response to stress and often involves the formation of a stress-resistant dormant form. The proximate and ultimate causes of the connection between stress and sex in facultatively sexual lineages are unclear. Because most forms of stress result in the overproduction of cellular reactive oxygen species (ROS), we address the hypothesis that this connection involves ROS and possibly reflects the ancestral role of sex as an adaptive response to the damaging effects of stress-induced ROS (i.e. oxidative stress) . Here, we report that two antioxidants inhibit sexual induction in a facultatively sexual species-the multicellular green alga, Volvox carteri. Furthermore, the nature of the sex response and the effect of an iron chelator on sexual induction are consistent with sex being a response to the DNA-damaging effects of ROS. In addition, we present preliminary data to suggest that sex, cell-cycle arrest and apoptosis are alternative responses to increased levels of oxidative stress.

 

Rudel D, Sommer RJ

The evolution of developmental mechanisms
DEV BIOL 264 (1): 15-37 DEC 1 2003

Abstract:
Over the past two to three decades, developmental biology has demonstrated that all multicellular organisms in the animal kingdom share many of the same molecular building blocks and many of the same regulatory genetic pathways. Yet we still do not understand how the various organisms use these molecules and pathways to assume all the forms we know today. Evolutionary developmental biology tackles this problem by comparing the development of one organism to another and comparing the genes involved and gene functions to understand what makes one organism different from another. In this review, we revisit a set of seven concepts defined by Lewis Wolpert (fate maps, asymmetric division. induction, competence, positional information, determination, and lateral inhibition) that describe the characters of many developmental systems and supplement them with three additional concepts (developmental genomics, genetic redundancy, and genetic networks). We will discuss examples of comparative developmental studies where these concepts have guided observations on the advent of a developmental novelty. Finally, we identify a set of evolutionary frameworks, such as developmental constraints, cooption, duplication, parallel and convergent evolution, and homoplasy, to adequately describe the evolutionary properties of developmental systems. (C) 2003 Elsevier Inc. All rights reserved.

Kato-Minoura T, Okumura M, Hirono M, et al.

A novel family of unconventional actins in volvocalean algae
J MOL EVOL 57 (5): 555-561 NOV 2003

Abstract:
The unicellular green alga Chlamydomonas reinhardtii has two actin genes, one encoding a conventional actin (90% amino acid identity with mammalian actin), the other a highly divergent actin (64% identity) named novel actin-like protein (NAP). To see whether the presence of conventional and unconventional actins in a single organism is unique to C. reinhardtii, we searched for genomic sequences related to the NAP sequence in several other species of volvocalean algae. Here we show that Chlamydomonas moewusii and Volvox carteri also have, in addition to a conventional actin, an unconventional actin similar to the C. reinhardtii NAP. Analyses of the deduced protein sequences indicated that the NAP homologues form a distinct group derived from conventional actin.

Golstein P, Aubry L, Levraud JP

Cell-death alternative model organisms: Why and which?
NAT REV MOL CELL BIO 4 (10): 798-807 OCT 2003

Abstract:
Classical model organisms have helped greatly in our understanding of cell death but, at the same time, night have constrained it. The use of other, nor-classical model organisms from all biological kingdoms could reveal undetected molecular pathways and better-defined morphological types of cell death. Here we discuss what is known and what might be learned from these alternative model systems.

Ohta H, Suzuki T, Ueno M, et al.

Extrinsic proteins of photosystem II - An intermediate member of the PsbQ protein family in red algal PSII
EUR J BIOCHEM 270 (20): 4156-4163 OCT 2003

Abstract:
The oxygen-evolving photosystem II (PS II) complex of red algae contains four extrinsic proteins of 12 kDa, 20 kDa, 33 kDa and cyt c-550, among which the 20 kDa protein is unique in that it is not found in other organisms. We cloned the gene for the 20-kDa protein from a red alga Cyanidium caldarium. The gene consists of a leader sequence which can be divided into two parts: one for transfer across the plastid envelope and the other for transfer into thylakoid lumen, indicating that the gene is encoded by the nuclear genome. The sequence of the mature 20-kDa protein has low but significant homology with the extrinsic 17-kDa (PsbQ) protein of PS II from green algae Volvox Carteri and Chlamydomonas reinhardtii, as well as the PsbQ protein of higher plants and PsbQ-like protein from cyanobacteria. Cross-reconstitution experiments with combinations of the extrinsic proteins and PS Its from the red alga Cy. caldarium and green alga
Ch. reinhardtii showed that the extrinsic 20-kDa protein was functional in place of the green algal 17-kDa protein on binding to the green algal PS II and restoration of oxygen evolution. From these results, we conclude that the 20-kDa protein is the ancestral form of the extrinsic 17-kDa protein in green algal and higher plant PS IIs. This provides an important clue to the evolution of the oxygen-evolving complex from prokaryotic cyanobacteria to eukaryotic higher plants. The gene coding for the extrinsic 20-kDa protein was named psbQ' (prime).

 

Cole DG, Reedy MV

Algal morphogenesis: How Volvox turns itself inside-out
CURR BIOL 13 (19): R770-R772
SEP 30 2003

Abstract:
During its development, the multicellular green alga Volvox undergoes inversion, in which spherical embryos turn their multicellular sheet completely inside out. A mutant analysis has revealed that a novel kinesin motor protein is essential for completing this process.

Voigt J, Frank R

14-3-3 proteins are constituents of the insoluble glycoprotein framework of the Chlamydomonas cell wall
PLANT CELL 15 (6): 1399-1413 JUN 2003

 

Abstract:
The cell wall of the unicellular green alga Chlamydomonas reinhardtii consists predominantly of Hyp-rich glycoproteins, which also occur in the extracellular matrix of multicellular green algae and higher plants. In addition to the Hyp-rich polypeptides, the insoluble glycoprotein framework of the Chlamydomonas cell wall contains minor amounts of
14-3-3 proteins, as revealed by immunochemical studies and mass spectroscopic analysis of tryptic peptides. Polypeptides immunologically related to the 14-3-3 proteins also were found in the culture medium of Chlamydomonas. The levels of two of these 14-3-3-related polypeptides were decreased in the culture medium of the wall-deficient mutant cw-15. These findings indicate that 14-3-3 proteins are involved in the cross-linking of Hyp-rich glycoproteins in the Chlamydomonas cell wall.

 

Michod RE, Nedelcu AM

On the reorganization of fitness during evolutionary transitions in individuality
INTEGR COMP BIOL 43 (1): 64-73 FEB 2003 (for pdf click here)

Abstract:
The basic problem in an evolutionary transition is to understand how a group of individuals becomes a new kind of individual, possessing the property of heritable variation in fitness at the new level of organization. During an evolutionary transition, for example, from single cells to multicellular organisms, the new higher-level evolutionary unit (multicellular organism) gains its emergent properties by virtue of the interactions among lower-level units (cells). We see the formation of cooperative interactions among lower-level units as a necessary step in evolutionary transitions; only cooperation transfers fitness from lower levels (costs to group members) to higher levels (benefits to the group). As cooperation creates new levels of fitness, it creates the opportunity for conflict between levels as deleterious mutants arise and spread within the group. Fundamental to the emergence of a new higher-level unit is the mediation of conflict among lower-level units in favor of the higher-level unit. The acquisition of heritable variation in fitness at the new level, via conflict mediation, requires the reorganization of the basic components of fitness (survival and reproduction) and life-properties (such as immortality and totipotency) as well as the co-option of lower-level processes for new functions at the higher level. The way in which the conflicts associated with the transition in individuality have been mediated, and fitness and general life-traits have been re-organized, can influence the potential for further evolution (i.e., evolvability) of the newly emerged evolutionary individual. We use the volvocalean green algal group as a model-system to understand evolutionary transitions in individuality and to apply and test the theoretical principles presented above. Lastly, we discuss how the different notions of individuality stem from the basic properties of fitness in a multilevel selection context.

 

Kirk DL

Seeking the ultimate and proximate causes of Volvox multicellularity and cellular differentiation
INTEGR COMP BIOL 43 (2): 247-253 APR 2003

Abstract:
 
Volvox and its relatives provide an exceptional model for integrative studies of the evolution of multicellularity and cellular differentiation. The volvocine algae range in complexity from unicellular Chlamydomonas through several colonial genera with a single cell type, to multicellular Volvox with its germsoma division of labor. Within the monophyletic family Volvocaceae, several species of Volvox have evolved independently in different lineages, the ultimate cause presumably being the advantage that large size and cellular differentiation provide in competing for limiting resources such as phosphorous. The proximate causes of this type of evolutionary transition are being studied in V carteri. All volvocine algae except Volvox exhibit biphasic development: cells grow during a motile, biflagellate phase, then they lose motility and divide repeatedly during the reproductive phase. In V carteri three kinds of genes transform this ancestral biphasic program into a dichotomous one that generates non-motile reproductive cells and biflagellate somatic cells with no reproductive potential: first the gls genes act in early embryos to cause asymmetric division and production of large-small sister-cell pairs; then lag genes act in the large cells to repress the biflagellate half of the ancestral program, while regA acts in the small cells to repress the reproductive half of the program. Molecular-genetic analysis of these genes is progressing, as will be illustrated with regA, which encodes a transcription factor that acts in somatic cells to repress nuclear genes encoding chloroplast proteins. Repression of chloroplast biogenesis prevents these obligately photoautotrophic cells from growing, and since they cannot grow, they cannot reproduce.

Mori T, Kuroiwa H, Higashiyama T, et al.

Identification of higher plant GlsA, a putative morphogenesis factor of gametic cells
BIOCHEM BIOPH RES CO 306 (2): 564-569
JUN 27 2003

Abstract:
GlsA has been identified in an asexual-reproductive-cell (gonidia)-deficient mutant of Volvox as a chaperone-like protein essential for gonidia production. In this study, we isolated an angiosperm glsA (LlglsA) gene expressed during Lilium longiflorum pollen development. Immunoblot analyses showed that the strong LlGlsA expression occurred in the generative cell and its pattern during pollen development corresponded to that of alpha-tubulin. Morphological analyses succeeded in visualizing the dispersion of the strong LlGlsA signal in developing generative cells. In addition, multiple-immunofluorescence staining of LIGNA and alpha-tubulin revealed that some of the dot-like LlGlsA signals were co-localized with microtubule filaments. From those results, we suggest that angiosperm GlsA functions as a chaperone modifying various structures during male gametic cell formation. (C) 2003 Elsevier Science (
USA). All rights reserved.

 

Nishii I, Ogihara S, Kirk DL

A kinesin, InvA, plays an essential role in Volvox morphogenesis
CELL 113 (6): 743-753
JUN 13 2003

Abstract:
In Volvox carted adults, reproductive cells called gonidia are enclosed within a spherical monolayer of biflagellate somatic cells. Embryos must "invert" (turn inside out) to achieve this configuration, however, because at the end of cleavage the gonidia are on the outside and the flagellar ends of all somatic cells point inward. Generation of a bend region adequate to turn the embryo inside out involves a dramatic change in cell shape, plus cell movements. Here, we cloned a gene called invA that is essential for inversion and found that it codes for a kinesin localized in the cytoplasmic bridges that link all cells to their neighbors. In invA null mutants, cells change shape normally, but are unable to move relative to the cytoplasmic bridges. A normal bend region cannot be formed and inversion stops. We conclude that the InvA kinesin provides the motile force that normally drives inversion to completion.

 

Bonner JT

On the origin of differentiation
J BIOSCIENCES 28 (4): 523-528 JUN 2003

 

Abstract:
Following the origin of multicellularity in many groups of primitive organisms there evolved more than one cell type. It has been assumed that this early differentiation is related to size - the larger the organism the more cell types. Here two very different kinds of organisms are considered: the volvocine algae that become multicellular by growth, and the cellular slime moulds that become multicellular by aggregation. In both cases there are species that have only one cell type and others that have two. It has been possible to show that there is a perfect correlation with size: the forms with two cell types are significantly larger-than those with one. Also in both groups there are forms of intermediate size that will vary from one to two cell types depending on the size of the individuals, suggesting a form of quorum sensing. These observations reinforce the view that size plays a critical role in influencing the degree of differentiation.

 

Michod RE, Nedelcu AM, Roze D

Cooperation and conflict in the evolution of individuality IV. Conflict mediation and evolvability in Volvox carteri
BIOSYSTEMS 69 (2-3): 95-114 MAY 2003 (for pdf click here)

Abstract:
The continued well being of evolutionary individuals (units of selection and evolution) depends upon their evolvability, that is their capacity to generate and evolve adaptations at their level of organization, as well as their longer term capacity for diversifying into more complex evolutionary forms. During a transition from a lower- to higher-level individual, such as the transition between unicellular and multicellular organisms, the evolvability of the lower-level (cells) must be restricted, while the evolvability of the new higher-level unit (multicellular organism) must be enhanced. For these reasons, understanding the factors leading to an evolutionary transition should help us to understand the factors underlying the emergence of evolvability of a new evolutionary unit. Cooperation among lower-level units is fundamental to the origin of new functions in the higher-level unit. Cooperation can produce a new more complex evolutionary unit, with the requisite properties of heritable fitness variations, because cooperation trades fitness from a lower-level (the costs of cooperation) to the higher-level (the benefits for the group). For this reason, the evolution of cooperative interactions helps us to understand the origin of new and higher-levels of fitness and organization. As cooperation creates a new level of fitness, it also creates the opportunity for conflict between levels of selection, as deleterious mutants with differing effects at the two levels arise and spread. This conflict can interfere with the evolvability of the higher-level unit, since the lower and higher-levels of selection will often "disagree" on what adaptations are most beneficial to their respective interests. Mediation of this conflict is essential to the emergence of the new evolutionary unit and to its continued evolvability. As an example, we consider the transition from unicellular to multicellular organisms and study the evolution of an early-sequestered germ-line in terms of its role in mediating conflict between the two levels of selection, the cell and the cell group. We apply our theoretical framework to the evolution of germ/soma differentiation in the green algal group Volvocales. In the most complex member of the group, Volvox carteri, the potential conflicts among lower-level cells as to the "right" to reproduce the higher-level individual (i.e. the colony) have been mediated by restricting immortality and totipotency to the germ-line. However, this mediation, and the evolution of an early segregated germ-line, was achieved by suppressing mitotic and differentiation capabilities in all post-embryonic cells. By handicapping the soma in this way, individuality is ensured, but the solution has affected the long-term evolvability of this lineage. We think that although conflict mediation is pivotal to the emergence of individuality at the higher-level, the way in which the mediation is achieved can greatly affect the longer-term evolvability of the lineage. (C) 2002 Elsevier Science Ireland Ltd. All rights reserved.

 

Laflamme M, Lee RW

Mitochondrial genome conformation among CW-group chlorophycean algae
J PHYCOL 39 (1): 213-220 FEB 2003

 

Abstract:
Most green algal taxa have circular-mapping mitochondrial genomes, whereas some have linear genome- or subgenomic-sized mitochondrial DNAs (mtDNA). It is not clear, however, if the circular-mapping genomes represent genome-sized circular molecules, if such circular molecules and the linear forms are the predominant in vivo mtDNA structures, or if the linear forms arose only once or multiple times among extant green algal lineages. We therefore examined the DNA components detected with homologous mtDNA probes after pulsed-field gel electrophoresis of total cellular DNA from the chlorophycean basal bodies displaced clockwise(CW)-group taxa Chlamydomonas reinhardtii and Chlamydomonas moewusii. For C. reinhardtii , the 15.8-kb linear mtDNA was the only DNA component detected, and there was no evidence of circular or large linear precursors of this DNA. In the case of C. moewusii , which is known to have a circular-mapping 22.9-kb mitochondrial genome, three DNA components were detected; these appeared to be circular (relaxed and supercoiled) and genome-sized linear DNA molecules, the latter of which likely resulted from random double-strand breaks in the circular forms during DNA isolation. In further studies, DNA from additional CW-group taxa was examined using conventional gel electrophoresis and DNA-filter blot analysis with C. reinhardtii and C. moewusii mtDNA probes. We conclude that all taxa from the "Volvox clade" (sensu Nakayama et al. 1996) of the CW-group have genome- or subgenomic-sized linear mtDNAs as their predominant mtDNA form and that these arose from a genome-sized circular form in an ancestor that existed near the base of this clade.

 

Hallmann A

Extracellular matrix and sex-inducing pheromone in Volvox
INT REV CYTOL 227: 131-+ 2003

 

Schmitt R, Sumper M

Developmental biology - How to turn inside out
NATURE 424 (6948): 499-500 JUL 31 2003

 

Desnitskiy AG.

Peculiarities of the geographical distribution of coenobial volvocine algae (Volvocaceae, Chlorophyta).

Botanical Journal (St Petersburg). 2003. Vol. 88. N 11. pp. 52-61.(in Russian, with English Summary)

 

Abstract:

Data on the geographical distribution of 36 species from 7 genera of the family Volvocaceae sensu Nozaki (Pandorina, Volvulina, Yamagishiella, Eudorina, Platydorina, Pleodorina and Volvox) have been summarized. Both cosmopolitan species and species with local distribution have been detected. An attempt was made to trace a correlation of latitudinal distribution of the coenobial volvocine algae with obligatory differentiated somatic cells (22 species of the genera Volvox and Pleodorina) with peculiarities of proceeding of the cell divisions during asexual developmental cycle. In high latitudes of the Northern Hemisphere (northward of 50-57° north) only 3 species of Volvox occur, in which the formation of new coenobia (a series of consecutive gonidial divisions) starts with the light period (in the morning), the rate of division is slow and the gonidial divisions are temporarily blocked in darkness.