Botanica Pacifica. A journal of plant science and conservation. 2015. 4(2): 59–70 DOI: 10.17581/bp.2015.04216 Phylogenesis, Origin and Kinship of the Charophytic Algae Galina­M.­Palamar­Mordvintseva­1,­Petro­M.­Tsarenko­1 *, Sophia Barinova 2 Galina­M.­Palamar­Mordvintseva­1 ABSTRACT e­mail:­[email protected]. We analyzed phylogenetic reconstructions of charophytes (streptophyte) algae Petro­M.­Tsarenko­1 * and assessed the importance of new phylogenetic hypotheses on some aspects e­mail:­[email protected] of the evolution of plants. We give attention to the progress made in modern Sophia Barinova 2 mo le cular biology study, and the occurrence of biological features and life cycles e­mail:­[email protected] of organisms and multicellular gravitropism based on the results of molecular phy sio lo gy and genome evolution. We analyzed generalized information on the sig­ni­fi­cance­and­place­of ­Сharophyta­in­the­development­of ­plants,­how­these­ 1 N.G. Kholodny Institute of Botany, plants were settled on land and their further development. We represent ideas NAS­of ­Ukraine,­Kiev,­Ukraine about the modern interpretation of ancestral forms and consanguinity of this group of algae with other taxa of plants. We used a phylogenetic approach in 2 Institute of Evolution, University of Haifa, Haifa, Israel ana ly sis of formation and methods of cell division, with a deeper concept of com ple xi ty of the process of plant organization (the transition from unicellular to­multicellular).­We­assessed­evolutionary­significance­of ­the­gravitropism­phe­ * corresponding author no menon and some features of molecular and physiological changes in plant or­ga­nisms­during­the­change­of ­environments­and­their­outlet­to­the­earth.­Sum­ Manuscript­received:­04.08.2015 ma rized and analyzed the changes in the modern paradigm in respect of group Review­completed:­02.10.2015 of streptophytes algae which can be assessed as a basis of other plants and close Accepted­for­publication:­10.10.2015 re­la­ted­to­Zygnematales­line­group­of ­algae­on­the­phylogenetic­tree. Published­online:­24.10.2015 Keywords: Сharophytes,­phylogenies,­evolution,­origin,­phylogenetic­relationships РЕЗЮМЕ Паламарь-Мордвинцева Г.М., Царенко П.М., Баринова С.С. Филоге- нез, происхождение и родственные связи харофитовых водорослей. Представлены­ материалы­ анализа­ филогенетических­ реконструкций­ харо­ фи­товых­водорослей,­а­также­освещена­значимость­новых­филогенети­чес­ких­ гипотез­о­некоторых­аспектах­эволюции­растений,­с­учетом­современ­ного­ прогресса­ в­ их­ молекулярно­биологическом­ изу­че­нии,­ раскрытии­ осо­бен­ ностей­биологии­жизненных­циклов­и­развития­ор­га­низмов,­воз­ник­но­ве­нии­ многоклеточности­и­гравитропизма,­с­учетом­ре­зуль­татов­молекулярной­фи­ зио­ло­гии­ и­ эволюции­генома.­ Обобщены­ дан­ные­ о­ формировании­ пред­ став­лений­о­значимости­харофитов­в­ста­нов­лении­мира­растений,­за­се­ле­нии­ ими­суши­и­дальнейшего­их­развития.­Обсуждается­современная­трак­тов­ка­ анцестральных­форм­и­родственных­связях­этой­группы­водорослей­с­дру­ гими­таксонами­рас­ти­тельного­мира.­С­использованием­филогенетическо­ го­подхода­показаны­особенности­формирования­и­деления­клетки,­а­так­же­ особенности­про­цесса­усложнения­растительной­организации­(переход­от­ од­но­кле­точ­ности­к­многоклеточности)­и­молекулярно­физиологичес­ких­из­ ме­нений­растительных­организмов­в­период­смены­среды­обитания­и­выхода­ их­на­сушу.­Обобщены­и­проанализированы­материалы­относительно­сме­ ны­или­модификации­современной­парадигмы­о­близкородст­вен­ной­группе­ стрептофитовых­ водорослей­ к­ базису­ других­ растений­ и­ род­ственной­ се­ стринской­линии­Zygnematales­на­филогенетическом­древе. Ключевые слова: харофиты,­филогенез,­эволюция,­родственные­связи Charophyte­algae­is­a­monophyletic­group­of ­strep­to­ Classic­morphological­and­taxonomic­ideas­of ­direct­fa­­ phyte lines in the evolution of plants (Viridiplantae), which mi­ly­con­nec­tions­of ­charophytes­algae­(Charales)­with­cryp­ dissociated­itself­from­chlorophyte­lines­more­than­450­mil­ to gams or the consideration of the group as direct ancestors li on years ago. This taxonomic group now unites uni cellular of ­"higher­plants"­have­not­been­confirmed­by­molecular­ and­ multicellular­ organisms­ (monoсytes­ and­ coenocytes)­ bio lo gi cal research. Also, there is no consensus about how with­different­morphological­structure­of ­thalluses,­are­cha­ these related groups of plants are phylogenetically related to rac terized by many morphological and cy to lo gical features other plants. In particular, if Mesostigma viride Lauterborn is si mi lar to other representatives of Strep to phyta, growing in the ancestral form of Viridiplantae (Bhattacharia & Medlin di verse environment such as water and soil. 1998,­ Lemieux­ et­ al.­ 2000,­ 2007,­ Rodriguez­Ezpeleta­ et­ ©Botanical Garden-Institute FEB RAS. 2015. Special issue “Plant evolution at levels from molecular to ecosystem” 59 Palamar-Mordvintseva et al. al.­2007)­on­the­base­of ­genomic­and­chloroplast­nuclear­ of genes, often from one to several cellular structures, or mi to chon d rial composition, or considered as complex of mul tiple groups of organisms.­As­a­result,­the­significant­ species M. viride and Chlorokybus atmophyticus­ Geitler­ (Le­ eviden ces, which support phylogenetic developments, were mieux­et­al.­2007),­then­the­sisterly­phylogenetic­lines­in­this­ ob­tained­ (Delsue­ et­ al.­ 2003,­ Qiu­ 2008,­ Becker­ &­ Marin­ case is deduced from different taxonomic groups of the 2009,­ Gontcharov­ 2009,­ Finet­ et­ al.­ 2010,­ Zong­ et­ al.­ Charophytes – Zyg ne ma ta les /Zygnematophyceae (Tur mel 2013).­ In­ addition,­ the­ use­ of ­ modern,­ highly­ developed­ et­al.­2006,­Gont­cha­rov­2008,­2009,­Wodniok­et­al.­2011,­ se­quen­cing­technology­and­increasing­the­number­of ­inves­ Timme­ et­ al.­ 2012,­ Zong­ et­ al.­ 2013,­ Ruhfel­ et­ al.­ 2014,­ ti gated taxa have improved our understanding in problem Wic­kett­et­al.­2014),­Coleochaetales­(Turmel,­Gagnon­et­al.­ sol ving of phylogenetic studies and enriched the base for 2009,­Turmel­et­al.­2009)­or­Coleochaetales­with­Zyg­ne­ma­ the­complicated­phylogenetic­reconstructions­(Brinkmann­ ta­les­(Finet­et­al.­2010,­Laurin­Lemay­et­al.­2012).­ &­Phillipe­2008).­In­the­traditional­taxonomy­we­can­use­ The­aim­of ­this­work­is­to­analyze­and­generalize­in­for­ all sources of newest phylogenetic data that give us much mation about the modern pro g ress of the Charophyte al gae more information and coverage of diverse approaches to phylogenetic­ reconstructions,­ coverage­ of ­ the­ signi­fi­cance­ decrypt historical models of evolution. of ­ new­ phylogenetic­ hypotheses­ on­ some­ key­ as­pects­ of ­ It­has­clearly­been­established­that­‘green­plants’,­Vi­ri­ the evolution of plants with special attention to mo le cular di­plan­tae­(Levis­&­McCourt­2004,­Qiu­2008,­Becker­&­Ma­ biological research, the disclosure of life cycle fea tures, the rin­2009,­Gontcharov­2008,­2009,­Finet­et­al.­2010),­unite­ causes of the emergence of multicellularity and gravitropism, all the ‘green’ algae with land plants (Embryophyte). They as well as to analyze the results and ideas of molecular are­ the­ monophyletic­ group­ of ­ organisms­ with­ an­ ex­t­ra­ physiology in evolution of the charophytes ge nomes. The or­dinary­ variety­ of ­ morphology,­ cell­ structure,­ life­ his­to­ tasks­ are­ associated­ with­ the­ preparation­ of ­ Charophyte­ ry, reproduction, and biochemistry. ‘Green plants’ has been algal­flora­of ­Ukraine­and­analyses­of ­va­ri­ous­aspects­that­ divided­ early­ enough­ into­ two­ evolutionary­ phyla­ (lines):­ affect this group of aquatic plants. Lo ca tion and value of the green­ algae­ (Chlorophyta)­ and­ charophytic­ algae­ (Cha­ro­ Charophyte algae in the organic world have been discussed phyta) with embryophyte (Streptophyta). This se pa ra tion earlier­(Palamar­Mordvintseva­&­Tsarenko­2009). happened­about­725–1,200­million­years­ago­(Hed­ges­et­al.­ 2004,­Yoon­et­al.­2004,­Zimmer­et­al.­2007).­In­com­pa­ri­ Phylogenesis of the Сharophytes son­with­the­chlorophyte­line­of ­evolution,­which­ac­com­ Reconstruction of the phylogeny of organisms was the mo dates most types of traditional division Chlo ro phyta main­aim­of ­biology­after­the­publication­of ­Darwin’s­the­o­ (100­ge­ne­ra­and­more­then­10000­species),­the­Cha­ro­phyte­ ry­of ­the­evolution­of ­organisms­(Darwin­1859).­The­emer­ (Strep to phyte) line contains relatively few existing green gence­of ­land­plants­(Embryophyte)­was­one­of ­the­grea­ cha­ro­phy­tic­ algae­ (about­ 65­ genera­ and­ several­ thousand­ test events in the history of living things, which caused an spe cies), which, together with almost half a mil li on species irreversible evolutionary process in the formation of life on of land plants (Embryophyte) compose the Strep tophyta Earth.­Setting­cladistics­principles­(Henning­1966)­and­scien­ di­vi­sion­(sensu­Bremer­et­al.­1987),­or­the­Cha­ro­phyta­(sen­ tific­achievements­of ­1960–1970­years­(Pickett­Heaps­1967,­ su­Levis­&­McCourt­2004). 1975,­Picket­Heaps­&­Marchant­1972,­Marchant­&­Pic­kett­ The descendants of the Charophyte algae occupied land Heaps­1973),­which­outlined­precise­conceptual­frame­works­ ha­bi­tats­about­450–470­million­years­ago­that­became­an­ex­ for the disclosure of relationships between or ga nisms and tra or dinary event in the evolution of life on Earth (Graham formal separation of charophytic algae in the modern sense 1993,­Kenrick­&­Crane­1997,­Bateman­et­al.­1998). of the group’s volume