MA H

GTP-BINDING PROTEINS IN PLANTS - NEW MEMBERS OF AN OLD FAMILY
PLANT MOL BIOL 26 (5): 1611-1636 DEC 1994

Abstract:
Regulatory guanine nucleotide-binding proteins (G proteins) have been studied extensively in animal and microbial organisms, and they are divided into the heterotrimeric and the small (monomeric) classes. Heterotrimeric G proteins are known to mediate signal responses in a variety of pathways in animals and simple eukaryotes, whiole small G proteins perform diverse functions including signal transduction, secretion, and regulation of cytoskeleton. In recent years, biochemical analyses have produced a large amount of information on the presence and possible functions of G proteins in plants. Further, molecular cloning has clearly demonstrated that plants have both heterotrimeric and small G proteins. Although the functions of the plant heterotrimeric G proteins are yet to be determined, expression analysis of an Arabidopsis Ga protein suggests that it may be involved in the regulation of cell division and differentiation. In contrast to the very few genes cloned thus far that encode heterotrimeric G proteins in plants, a large number of small G proteins have been identified by molecular cloning from various plants. In addition, several plant small G proteins have been shown to be functional homologues of their counterparts in animals and yeasts. Future studies using a number of approaches are likely to yield insights into the role plant G proteins play.

 

VALIN P, GOULARD B, SANIELEVICI M

TOPOLOGY DEPENDENCE IN LATTICE SIMULATIONS OF NONLINEAR PDES ON A MIMD COMPUTER
INT J MOD PHYS C 5 (6): 957-971 DEC 1994

 

Abstract:
We tested the parallelization of explicit schemes for the solution of non-linear classical field theories of complex scalar fields which are capable of simulating; hadronic collisions. Our attention focused on collisions in a fractional model with a particularly rich inelastic spectrum of final states. Relativistic collisions of all types were performed by computer on large lattices (64 to 256 sites per dimension). The stability and accuracy of the objects were tested by the use of two other methods of solutions: Pseudo-spectral and semi-implicit. Parallelization of the Fortran code on a 64-transputer MIMD Volvox machine revealed, for certain topologies, communication deadlock and less-than-optimum routing strategies when the number of transputers used was less than the maximum. The observed speedup, for N transputers in an appropriate topology, is shown to scale approximately as N, but the overall gain in execution speed, for physically interesting problems, is a modest 2-3 when compared to state-of-the-art workstations.

 

SCHULZE E, NAGEL S, GAVENIS K, et al.

STRUCTURALLY DIVERGENT HISTONE H1 VARIANTS IN CHROMOSOMES CONTAINING HIGHLY CONDENSED INTERPHASE CHROMATIN
J CELL BIOL 127 (6): 1789-1798 Part 2 DEC 1994

Abstract:
Condensed and late-replicating interphase chromatin in the Dipertan insect Chironomus contains a divergent type of histone H1 with an inserted KAPKAP repeat that is conserved in single H1 variants of Caenorhabditis elegans and Volvox carteri. H1 peptides comprising the insertion interact specifically with DNA. The Chironomid Glyptotendipes exhibits a corresponding correlation between the presence of condensed chromosome sections and the appearance of a divergent H1 subtype. The centromere regions and other sections of Glyptotendipes barbipes chromosomes are inaccessible to immunodecoration by anti-H2B and anti-H1 antibodies one of which is known to recognize nine different epitopes in all domains of the H1 molecule. Microelectrophoresis of the histones from manually isolated unfixed centromeres revealed the presence of H1 and core histones. H1 genes of G. barpipes were sequenced and found to belong to two groups. H1 II and H1 III are rather similar but differ remarkably from H1 I. About 30% of the deduced amino acid residues were found to be unique to H1 I. Most conspicuous is the insertion, SPAKSPGR, in H1 I that is lacking in H1 II and H1 III and at its position gives rise to the sequence repeat SPAKSPAKSPGR. The homologous H1 I gene in Glyptotendipes salinus encodes the very similar repeat TPAKSPAKSPGR. Both sequences are structurally related to the KAPKAP repeat in H1 I-1 specific for condensed chromosome sites in Chironomus and to the SPKKSPKK repeat in sea urchin sperm H1, lie at almost the same distance from the central globular domain, and could interact with linker DNA in packaging condensed chromatin.

 

WOESSNER JP, MOLENDIJK AJ, VANEGMOND P, et al.

DOMAIN CONSERVATION IN SEVERAL VOLVOCALEAN CELL-WALL PROTEINS
PLANT MOL BIOL 26 (3): 947-960 NOV 1994

Abstract:
Based on our previous work demonstrating that (SerPro)(x) epitopes are common to extensin-like cell wall proteins in Chlamydomonas reinhardtii, we looked for similar proteins in the distantly related species C. euganzetos. Using a polyclonal antiserum against a (SerPro)(10) oligopeptide, we found distinct sets of stage-specific polypeptides immunoprecipitated from in vitro translations of C. eugametos RNA. Screening of a C. eugametos cDNA expression library with the antiserum led to the isolation of a cDNA (WP6) encoding a (SerPro)(x)-rich multidomain wall protein. Analysis of a similarly selected cDNA (VSP-3) from a C. reinhardtii cDNA expression library revealed that it also coded for a (SerPro)(x)-rich multidomain wall protein. The C-terminal rod domains of VSP-3 and WP6 are highly homologous, while the N-terminal domains are dissimilar; however, the N-terminal domain of VSP-3 is homologous to the globular domain of a cell wall protein from Volvox carteri. Exon shuffling might be responsible for this example of domain conservation over 350 million years of volvocalean cell wall protein evolution.

 

 

DIETMAIER W, FABRY S

ANALYSIS OF THE INTRONS IN GENES ENCODING SMALL G-PROTEINS
CURR GENET 26 (5-6): 497-505 NOV-DEC 19

 

Abstract:
Because all small G proteins (SGPs) possess a very similar array of structural and functional domains, they are obvious candidates for examining the relationships postulated to exist between the exon-intron structure of genes and the domain structure of the encoded proteins. To address this issue, and to possibly gain insight into the evolution of their introns, we have analyzed positions, sizes, and sequences of 125 introns from 28 SGP genes. These introns were found to be distributed in 60 different locations throughout the aligned sequences, with a preference for the 5'-half of the genes. More than 50% of the positions were found to be shared by two or more genes, and genes encoding SGPs of very similar amino acid sequence (i.e., isotypes) in quite closely related species tend to have most, or all, of their introns in identical locations, indicating a common evolutionary origin (homologous introns). However, with few exceptions, no statistically significant sequence similarity or common folding motif was found between homologous intron pairs. Only three intron positions are shared between members of distantly related SGP subfamilies. These three potentially ancient intron locations fall between regions encoding alpha-helices or beta-sheets, but two of them interrupt regions encoding known functional (guanosine-nucleotide-binding) modules. Intron positions that are occupied only in single genes, or in genes encoding very similar SGPs, do not show any preferential distribution with respect to regions encoding structural or functional motifs. This discordance between exon modules and structural and/or functional protein domains suggests that most, if not all, introns in modern SGP genes arose by independent insertion events after diversification of the various SGP subfamilies, and therefore probably did not participate in the early evolution of these genes.

 

HALLMANN A, SUMPER M

REPORTER GENES AND HIGHLY REGULATED PROMOTERS AS TOOLS FOR TRANSFORMATION EXPERIMENTS IN VOLVOX-CARTERI
P NATL ACAD SCI USA 91 (24): 11562-11566 NOV 22 1994

Abstract:
The multicellular alga Volvox is an attractive model for the study of developmental processes. With the recent report of successful transformation, regulated promoters as well as reporter genes working in this organism are now required. The Volvox genes encoding arylsulfatase and the extracellular glycoprotein ISG are strictly regulated. The former is transcribed only under conditions of sulfur starvation, whereas the latter operates under extreme developmental control-i.e., it is transcribed for only a few minutes in Volvox embryos at the stage of embryonic inversion. The gene encoding the sexual pheromone of Volvox carteri was placed under the control of the arylsulfatase promoter. In response to sulfur deprivation, V. carteri transformed by this construct synthesized and secreted biologically active pheromone. In addition, the gene encoding Volvox arylsulfatase was placed under the control of the ISG promoter. Transformed algae synthesized arylsulfatase mRNA only during embryonic inversion. These experiments demonstrate the usefulness of both the arylsulfatase and the sexual.

 

WOESSNER JP, GOODENOUGH UW

VOLVOCINE CELL-WALLS AND THEIR CONSTITUENT GLYCOPROTEINS - AN EVOLUTIONARY PERSPECTIVE
PROTOPLASMA 181 (1-4): 245-258 1994

 

Abstract:
Similarities in the composition of the extracellular matrix suggest that only some species of the unicellular Chlamydomonas are closely related to the colonial and multicellular flagellated members of the family Volvocaceae. The cell walls from all of the algae in this volvocine group contain a crystalline layer. This lattice structure can be used as a phylogenetic marker to divide Chlamydomonas species into distinct classes, only one of which includes the volvocacean algae. Similarly, not all species of Chlamydomonas are sensitive to each other's cell wall lytic enzymes, implying divergence of the enzyme's inner wall substrate. Interspecific reconstitution of the crystalline layer is possible between C. reinhardtii and the multicellular Volvox carteri, but not between C. reinhardtii and C. eugametos. The hydroxyproline-rich glycoproteins (HRGPs) which make up the crystalline layer in genera which have a similar crystal structure exhibit many homologies. Interestingly, the evolutionarily distant cell walls of C. reinhardtii and C. eugametos also contain some HRGPs displaying a few morphological and amino acid sequence homologies. The morphological similarities between the flagellar agglutinins (HRGPs responsible for sexual recognition and adhesion during the mating reaction) and the cell wall HRGPs leads to the proposal of a superfamily from which novel HRGPs (designed for self-assembly/recognition) can constantly evolve. Just as Variations in the wall HRGPs can lead to unique wall structures, new agglutinins facilitate sexual isolation of new species. Thus, the HRGPs could emerge as Valuable phylogenetic markers.

 

HUBER O, SUMPER M

ALGAL-CAMS - ISOFORMS OF A CELL-ADHESION MOLECULE IN EMBRYOS OF THE ALGA VOLVOX WITH HOMOLOGY TO DROSOPHILA FASCICLIN-I
EMBO J 13 (18): 4212-4222 SEP 15 1994

Abstract:
Proof that plants possess homologs of animal adhesion proteins is lacking. In this paper we describe the generation of monoclonal antibodies that interfere with cell-cell contacts in the 4-cell embryo of the multicellular alga Volvox carteri, resulting in a hole between the cells. The number of following cell divisions is reduced and the cell division pattern is altered drastically. Antibodies given at a later stage of embryogenesis specifically inhibit inversion of the embryo, a morphogenetic movement that turns the embryo inside out. Immunofluorescence microscopy localizes the antigen (Algal-CAM) at cell contact sites of the del eloping embryo. Algal-CAR I is a protein with a three-domain structure: an N-terminal extensin-like domain characteristic for plant cell walls and two repeats with homology to fasciclin I, a cell adhesion molecule involved in the neuronal development of Drosophila. Alternatively spliced variants of Algal-CAM mRNA were detected that are produced under developmental control. Thus, Algal-CAM is the first plant homolog of animal adhesion proteins.

 

NASS N, MOKA R, JAENICKE L

ADENYLYL-CYCLASE FROM THE GREEN-ALGA VOLVOX-CARTERI F NAGARIENSIS - PARTIAL-PURIFICATION AND CHARACTERIZATION
AUST J PLANT PHYSIOL 21 (5): 613-622 1994

Abstract:
In order to understand changes in cyclic adenylate levels of Volvox carteri during the process of sexual induction, we investigated the biochemical properties of its membrane-bound adenylyl cyclase. Membrane preparations possess low levels of Mg2+-dependent or Mn2+-dependent adenylyl cyclase activity. This activity was solubilised and then purified 7800-fold. The enzyme detergent complex has an apparent molecular mass of 100 kDa. Purified preparations contain a major ATP-binding protein of 33 kDa as shown by affinity labelling. The Mg2+-dependent basal enzyme activity is regulated by Ca2+, and is highest in the presence of 10(-7) M Ca2+, but is inhibited by Ca2+ above 10(-5) M. La3+ at 10(-4) M also blocks activity. Neither calmodulin nor its antagonists affect the enzyme activity, nor do the purified preparations interact with immobilised calmodulin. Further mediators of G-protein action (NaF, or GTP and its derivatives) and forskolins have no influence on the basal activity of this plant enzyme. The function of adenylyl cyclase in sexual induction of Volvox is discussed.

 

HOOPS HJ, LONG JJ, HILE ES

FLAGELLAR APPARATUS STRUCTURE IS SIMILAR BUT NOT IDENTICAL IN VOLVULINA-STEINII, EUDORINA-ELEGANS, AND PLEODORINA-ILLINOISENSIS (CHLOROPHYTA) - IMPLICATIONS FOR THE VOLVOCINE EVOLUTIONARY LINEAGE
J PHYCOL 30 (4): 679-689 AUG 1994

Abstract:
The colonial and multicellular members of the Volvocales can be arranged in order of increasing size and complexity as the ''volvocine series.'' This series is often assumed to reflect an evolutionary progression. The flagellar apparatuses of previously examined algae are not consistent with a simple lineage. The flagellar apparatuses of Astrephomene gubernaculifera Pocock, Gonium pectorale Muller, Platydorina caudata Kofoid, Volvox rousseletii G. S. West, and V. carteri f. weismannia (Powers) Iyengar differ from one another, and there is no apparent progression in flagellar apparatus features from the simple to complex colonial forms. We examined the flagellar apparatuses of Volvulina steinii Playfair, Eudorina elegans Ehr., and Pleodorina illinoisensis Kofoid and found then to be similar to one another. The basal bodies ave connected by a distal fiber that is offset to the anti side of the cell. Two microtubular rootlets originate on the inside of the basal bodies and extend toward the syn side. The other two rootlets are oriented perpendicular to the first two and are anti-parallel to each other. A coarsely striated component underlies the four-membered rootlets and extends to the basal bodies. A proximal fiber complex connects the two basal bodies. This complex consists of a branched striated component on the cis side of each basal body. One part extends toward the anti side of the cell, while the other extends into a fibrous component that runs between basal bodies. An additional structure extends in the anti direction from the trans side of each basal body. A fibrous component extends past one basal body in all four species. This component goes past the trans basal body in Volvulina steinii and the cis basal body in E. elegans and P. illinoisensis. The flagellar apparatuses of these organisms are similar to those of G. pectorale and Volvox carteri but different from the other colonial volvocalean algae examined. The algae examined in this study plus G. pectorale and V. carteri probably share a common evolutionary history that postdates the transition from the unicellular to colonial habit. Such a shared evolutionary history is a requirement of the volvocine hypothesis. However, we have not observed progressive changes in the flagellar apparatus correlated with increasing cell number, differentiation, and sexual specialization. Thus, it is possible, but not certain, that G. pectorale, Volvulina steinii, E. elegans, P. illinoisensis, and Volvox carteri may form part of a volvocine lineage.

 

KIRK DL

GERM-CELL SPECIFICATION IN VOLVOX-CARTERI
CIBA F SYMP 182: 2-15 1994

 

Abstract:
Volvox carteri illustrates with diagrammatic clarity Weismann's concept of an immortal germline that produces a mortal soma that will carry it for a time, but then perish. Each V. carteri adult consists of about 16 asexual reproductive cells (gonidia) in the interior of a sphere that consists at its surface of about 2000 biflagellate somatic cells. When mature, each gonidium divides to form a juvenile with this same cellular composition. Half-way through their maturation, juveniles hatch out of the parenteral spheroid, whereupon parental somatic cells undergo programmed death while juvenile gonidia prepare for a new round of reproduction. The first visible step in V. carteri germ-soma differentiation is asymmetric cleavage, which sets apart large gonidial initials from small somatic initials. Experimental analysis indicates that it is a difference in size, not any difference in cytoplasmic quality, that determines whether a cell will become germinal or somatic. Mutational and molecular studies lead to the following model for the genetic control of the germ-soma dichotomy: first, the gls locus acts to cause asymmetric division; then large cells activate a set of lag loci that suppress expression of somatic genes, while small cells activate the regA locus that suppresses gonidial genes.

 

SCHIEDLMEIER B, SCHMITT R, MULLER W, et al.

NUCLEAR TRANSFORMATION OF VOLVOX-CARTERI
P NATL ACAD SCI USA 91 (11): 5080-5084 MAY 24 1994

Abstract:
Stable nuclear transformation of Volvox carteri was achieved using the cloned V. carteri nitA(+) gene (which encodes nitrate reductase) to complement a nitA(-) mutation. Following bombardment of mutant cells with plasmid-coated gold particles, putative transformants able to utilize nitrate as a nitrogen source were recovered with an efficiency of approximate to 2.5 x 10(-5). DNA analysis indicated that the plasmid integrated into the genome, often in multiple copies, at sites other than the nitA locus. Cotransformants were recovered with a frequency of 40-80% when cells were cobombarded with a selected and an unselected marker. Thus, V. carteri becomes one of the simplest multicellular organisms that is accessible to detailed molecular studies of genes regulating cellular differentiation and morphogenesis.

 

KOUFOPANOU V

THE EVOLUTION OF SOMA IN THE VOLVOCALES
AM NAT 143 (5): 907-931 MAY 1994

 

Abstract:
The presence of soma and the manner in which it segregates from the germ line is a fundamental aspect of development. This article examines the origin and evolution of soma in the Volvocales, the flagellated forms of green algae by analyzing data on cell division and development of 370 species. Phylogenetic analysis suggests that the cell wall is a preadaptation for the evolution of large multicellular colonies with deterministic development. The allometry of soma and germ supports the idea that soma functions to provide functional flagella during the embryogenesis of large colonies (Volvocaceae). The need for soma arises from a constraint that prevents simultaneous flagellation and cell division of cells surrounded by rigid walls: basal bodies cannot remain attached to their flagella while migrating to the mitotic poles. The con-straint is different from those that may cause the separation of flagellation and cell division in metazoan cells. Apart from a few developmental variations, which may represent adaptations to larger size, the Volvocaceae can be obtained by heterochronic changes in the timing of cell division. Their small size compared to the size of nonflagellated relatives can be attributed to their locomotion by flagella, which limits the maximum amount of germ that can be carried by the soma. This limitation is manifested in a negative correlation between size and number of germ cells among the largest species of Volvocaceae (Volvox). the opposite of a positive one among the smaller species.

 

NOZAKI H, ITOH M

PHYLOGENETIC-RELATIONSHIPS WITHIN THE COLONIAL VOLVOCALES (CHLOROPHYTA) INFERRED FROM CLADISTIC-ANALYSIS BASED ON MORPHOLOGICAL DATA
J PHYCOL 30 (2): 353-365 APR 1994

Abstract:

 A cladistic analysis was used to deduce the phylogenetic relationships within the colonial Volvocales. Forty-one pairs of characters related to gross morphology and ultrastructure of vegetative colonies as well as asexual and sexual reproduction were analyzed based on parsimony, using the PAUP 3.0 computer program, for 25 species belonging to nine volvocacean and goniacean genera of the colonial Volvocales. Chlamydomonas reinhardtii Dangeard was the outgroup. The strict consensus tree indicated the presence of two monophyletic groups, one composed of all the volvocacean species analyzed in this study and the other containing the goniacean species except for the four-celled species Gonium sociale (Dujardin) Warming. In addition, these two groups constitute a large monophyletic group, to which G. sociale is a sister group. A new combination Tetrabaena socialis (Dujardin) Nozaki et Itoh and a new family Tetrabaenaceae Nozaki et Itoh are thus proposed for G. sociale. In addition, the analysis suggests that the volvocacean genera Eudorina and Pleodorina are paraphyletic groups, respectively, and that the monotypic genus Yamagishiella has no autapomorphic characters and represents primitive features of the anisogamous and oogamous genera of the Volvocaceae. Phylogenetic relationships within the Volvocaceae and the Goniaceae, as well as the various modes of sexual reproduction exhibited by these organisms, are discussed on the basis of the analysis.

 

HALLMANN A, SUMPER M

AN INDUCIBLE ARYLSULFATASE OF VOLVOX-CARTERI WITH PROPERTIES SUITABLE FOR A REPORTER-GENE SYSTEM - PURIFICATION, CHARACTERIZATION AND MOLECULAR-CLONING
EUR J BIOCHEM 221 (1): 143-150 APR 1 1994


Abstract:
The multicellular green flagellate Volvox carteri synthesizes a periplasmic arylsulfatase in response to sulfur deprivation. The inducible enzyme has been purified to homogeneity and characterized. The corresponding gene and cDNA have been cloned. Determination of the sequence of genomic clones and comparisons to the cDNA sequence, revealed sixteen introns and seventeen a exons that encode a 649-amino-acid polypeptide chain. Since the arylsulfatase enzyme is readily assayed using chromogenic substrates, but is not detectable in cells grown in sulfate-containing medium, the gene encoding arylsulfatase may be useful as a reporter gene in V. carteri. In addition, the highly regulated promoter of the arylsulfatase gene suggests its suitability as a tool for producing inducible expression vectors for cloned genes.

 

SEKIMOTO H, SONE Y, FUJII T

REGULATION OF EXPRESSION OF THE GENES FOR A SEX-PHEROMONE BY AN INDUCER OF THE SEX-PHEROMONE IN THE CLOSTERIUM PERACEROSUM-STRIGOSUM-LITTORALE COMPLEX
PLANTA 193 (1): 137-144 MAR 1994

Abstract:
A sex pheromone, protoplast-release-inducing protein (PR-IP), of the Closterium peracerosum-strigosum-littorale complex is known to induce the release of protoplasts from mating-type minus (Mt(-)) cells during sexual reproduction and to have two subunit polypeptides of 19 and 42 kDa. Here, we describe the regulation mechanism for the release of the PR-IP. The sex pheromone was fractionated to yield subunits of 19 and 42 kDa, respectively, and each subunit was treated with V8 protease and with CNBr. By reference to the partial amino-acid sequences of the digested polypeptides, oligo nucleotides were synthesized and used as primers for the combined reverse transcription-polymerase chain reaction. Amplified fragments of DNA, of 130 bp in the case of the 19-kDa subunit and of 330 bp in the case of the 42-kDa subunit, were obtained, sequenced, and used as probes to identify the respective transcripts. From the results of Northern hybridization, the sizes of transcripts were estimated to be 1.2 kb for the 18-kDa subunit and 1.4 kb for the 42-kDa subunit. These transcripts appeared transiently only when mating-type plus (mt(+)) cells were treated with another sex pheromone (PR-IP Inducer) for more than 4h in the light. By immunoblotting with anti-42-kDa-subunit antiserum, it was shown that PR-IP accumulated gradually in the medium but not in the mt(+) cells after treatment with PR-IP Inducer in the light. We suggest that PR-IP is synthesized de novo and secreted from mt(+) cells only after the perception of PR-IP Inducer that has been released from mt(-) cells in the light during the sexual reproduction of Closterium, An analysis by genomic Southern hybridization revealed that probes for the 19-kDa and 42-kDa peptides hybridized to 6.8-kb and 5.1-kb DNA fragments, respectively, after the digestion of the genome with EcoRI. These hybridized DNA fragments were obtained not only from the genome of mt(+) cells but also from the genome of mt(-) cells, in which no transcripts for PR-IP could be detected by Northern hybridization. On the basis of these results, we discuss the possibility that the expression of the gene for the two subunits of PR-IP might be critically dependent upon the action of putative sex-determining genes.

 

SCHIEDLMEIER B, SCHMITT R

REPETITIOUS STRUCTURE AND TRANSCRIPTION CONTROL OF A POLYUBIQUITIN GENE IN VOLVOX-CARTERI
CURR GENET 25 (2): 169-177 FEB 1994

 

Abstract:
Southern analysis indicated the presence of at least four ubiquitin gene loci in the Volvox carteri genome. Three of these, a polyubiquitin gene described here and a non-segregating ubiquitin gene pair, were assigned to two different linkage groups by RFLP mapping; the non-polymorphic fourth gene locus remained unassigned. The polyubiquitin gene was cloned and its 2,116-bp sequence determined. It contains six exons each interrupted by an intron at Gly35, and it encodes a pentameric polyubiquitin polypeptide consisting of five runs of 76 identical amino-acid residues and a C-terminal extension of one leucine. The five tandem repeats of coding units plus introns exhibit an unusually high degree of overall sequence identity indicating an efficient process of gene homogenization in this region of the V. carteri genome. S1 mapping revealed two closely-spaced transcription starts, 24 and 28 nucleotides downstream from a putative TATA sequence. Preceding the TATA box are two 14-bp conserved heat-shock elements (HSEs) and two octameric sequences closely resembling an yeast HSE. Consistent with a 1.6-kb transcript seen on Northern blots are two polyadenylation signals (TGTAA) located 99 bp and 169 bp downstream from the TGA translational stop. The polyubiquitin gene was transcribed throughout the Volvox life cycle with peaks in the 1.6-kb mRNA levels during pre-cleavage, cleavage, and post-inversion. In contrast, an 0.6-kb monoubiquitin transcript was abundant only at the pre-cleavage stage suggesting a different type of gene control. Heat shock increased the level of polyubiquitin mRNA, whereas the level of monoubiquitin mRNA was down-regulated.