Embryological features of Tofieldia glutinosa and their bearing on the early diversification of monocotyledonous plants

Background and Aims

Although much is known about the vegetative traits associated with early monocot evolution, less is known about the reproductive features of early monocotyledonous lineages. A study was made of the embryology of Tofieldia glutinosa, a member of an early divergent monocot clade (Tofieldiaceae), and aspects of its development were compared with the development of other early divergent monocots in order to gain insight into defining reproductive features of early monocots.

Methods

Field-collected developing gynoecial tissues of Tofieldia glutinosa were prepared for histological examination. Over 600 ovules were sectioned and studied using brightfield, differential interference contrast, and fluorescence microscopy. High-resolution digital imaging was used to document important stages of megasporogenesis, megagametogenesis and early endosperm development.

Key Results

Development of the female gametophyte in T. glutinosa is of a modified Polygonum-type. At maturity the female gametophyte is seven-celled and 11-nucleate with a standard three-celled egg apparatus, a binucleate central cell (where ultimately, the two polar nuclei will fuse into a diploid secondary nucleus) and three binucleate antipodal cells. The antipodal nuclei persist past fertilization, and the process of double fertilization appears to yield a diploid zygote and triploid primary endosperm cell, as is characteristic of plants with Polygonum-type female gametophytes. Endosperm development is helobial, and free-nuclear growth initially proceeds at equal rates in both the micropylar and chalazal endosperm chambers.

Conclusions

The analysis suggests that the shared common ancestor of monocots possessed persistent and proliferating antipodals similar to those found in T. glutinosa and other early-divergent monocots (e.g. Acorus and members of the Araceae). Helobial endosperm among monocots evolved once in the common ancestor of all monocots excluding Acorus. Thus, the analysis further suggests that helobial endosperm in monocots is homoplasious with those helobial endosperms that are present in water lilies and eudicot angiosperms.Key words: Tofieldia, Tofieldiaceae, Alismatales, monocots, embryology, female gametophyte, antipodals, development, endosperm     

Embryology of <Emphasis Type="Italic">Japonolirion</Emphasis> (Petrosaviaceae,Petrosaviales): a comparison with other monocots

Japonolirion osense, the sole species of the genus, endemic to Japan, which is placed together with Petrosavia in the Petrosaviaceae and the order Petrosaviales, is still poorly known with respect to systematic characters. Here I present an embryological study of the anther, ovule, and seed of J. osense. Japonolirion is characterized by a glandular anther tapetum, simultaneous cytokinesis in the microspore mother cell, two-celled mature pollen grains, anatropous and crassinucellate ovules, a two-cell-layered nucellar cap formed early in ovule development, antipodal cells hypertrophied in post-fertilization stages, the ab initio cellular mode of endosperm formation, and exotegmic seeds. Comparisons with the basal monocots Acorus (Acorales) and Araceae (Alismatales), and with the more derived monocots Nartheciaceae (Dioscoreales) and Velloziaceae/Triuridaceae (Pandanales), showed that Japonolirion is clearly distinct from those basal and more derived monocots, supporting a distinct position for Petrosaviaceae or Petrosaviales within the monocots. Extensive comparisons further suggest that the two-cell-layered nucellar cap, whose cells are rich in cytoplasm at the time of fertilization in Japonolirion and thus obviously function as the obturator, is likely to be a common characteristic of the basal monocots and may even be a link with the magnoliids.     

Gynoecium diversity and systematics in basal monocots

Gynoecium and ovule structure was comparatively studied in representatives of the basal monocots, including Acorales (Acoraceae), Alismatales (Araceae, Alismataceae, Aponogetonaceae, Butomaceae, Hydrocharitaceae, Junc‐aginaceae, Limnocharitaceae, Potamogetonaceae, Scheuchzeriaceae, Tofieldiaceae), Dioscoreales (Dioscoreaceae, Taccaceae), and Triuridaceae as a family of uncertain position in monocots. In all taxa studied the carpels or gynoecia are closed at anthesis. This closure is attained in different ways: (1) by secretion without postgenital fusion (Araceae, Hydrocharitaceae); (2) by partly postgenitally fused periphery but with a completely unfused canal (Alismataceae, Aponogetonaceae, Butomaceae, Limnocharitaceae, Scheuchzeriaceae, Dioscoreaceae, Taccaceae); (3) by completely postgenitally fused periphery but with an unfused canal in the centre (Acoraceae, Tofieldiaceae); (4) by complete postgenital fusion and without an (unfused) canal (Juncaginaceae, Potamogetonaceae). In many Alismatales (but without Araceae) carpels have two lateral lobes. The stigmatic surface is restricted to the uppermost part of the ventral slit (if the carpel is plicate); it is never distinctly double‐crested (Butomaceae?). Stigmas are commonly unicellular‐papillate and secretory in most taxa. The locules are filled with a (often) mucilaginous secretion in a number of taxa. Superficial (probably intrusive) ethereal oil cells were found in the carpel wall of Acorus gramineus (as in Piperales!). Idioblasts in carpels are otherwise rare. A number of basal monocots has orthotropous ovules, which is perhaps the plesiomorphic condition in the group. The presence of almost tenuinucellar (pseudocrassinucellar) ovules is relatively common (Acoraceae, many Araceae, some Alismatales s.s.), whereas completely tenuinucellar ovules are rare and do not characterize larger groups. However, crassinucellar ovules occur in the largest number of families among the study group (basal Araceae, many Alismatales s.s.) The outer integument is always annular in orthotropous ovules. The inner integument is often lobed and it mostly forms the micropyle, whereas the outer integument is always unlobed. Gynoecium structure supports the isolated position of Acoraceae as sister to all other monocots. However, in an overall view, if compared with all other families, Acoraceae clearly shows the greatest similarities with Araceae.     

Evolution of the monocot gynoecium: evidence from comparative morphology and development in Tofieldia, Japonolirion, Petrosavia and Narthecium

We present new comparative morphological and developmental data on gynoecia of three genera of early-divergent monocots: Tofieldia (Tofieldiaceae, Alismatales), Petrosavia and Japonolirion (Petrosaviaceae, Petrosaviales) and one lilioid monocot: Narthecium (Nartheciaceae, Dioscoreales). Our data show significant differences between the genera examined, and are congruent with the splitting of former Nartheciaceae sensu Tamura (1998) into families Tofieldiaceae, Petrosaviaceae NB-cosistent with later and Nartheciacae (APG II 2003). Our investigation confirms the presence of at least partial carpel fusion in all taxa examined. Previous data indicating apocarpy in Japonolirion, some Petrosavia and Tofieldia could be due to late postgenital carpel fusion in these plants. Syncarpy also characterises other early-divergent monocot lineages such as Acoraceae and Araceae. It is most parsimonious to regard syncarpy as a primitive condition for monocots, but an alternative scenario suggests that apocarpy is plesiomorphic among monocots, involving multiple origins of syncarpy. The latter hypothesis is supported by significant differences between gynoecia of early-divergent monocots, including different modes of carpel fusion.     

Plastid phylogenomics and molecular evolution of Alismatales

Past phylogenetic studies of the monocot order Alismatales left several higher‐order relationships unresolved. We addressed these uncertainties using a nearly complete genus‐level sampling of whole plastid genomes (gene sets representing 83 protein‐coding and ribosomal genes) from members of the core alismatid families, Tofieldiaceae and additional taxa (Araceae and other angiosperms). Parsimony and likelihood analyses inferred generally highly congruent phylogenetic relationships within the order, and several alternative likelihood partitioning schemes had little impact on patterns of clade support. All families with multiple genera were resolved as monophyletic, and we inferred strong bootstrap support for most inter‐ and intrafamilial relationships. The precise placement of Tofieldiaceae in the order was not well supported. Although most analyses inferred Tofieldiaceae to be the sister‐group of the rest of the order, one likelihood analysis indicated a contrasting Araceae‐sister arrangement. Acorus (Acorales) was not supported as a member of the order. We also investigated the molecular evolution of plastid NADH dehydrogenase, a large enzymatic complex that may play a role in photooxidative stress responses. Ancestral‐state reconstructions support four convergent losses of a functional NADH dehydrogenase complex in Alismatales, including a single loss in Tofieldiaceae.     

单子叶植物高级分类阶元系统演化: matK、rbcL和18S rDNA序列的证据

基于两个叶绿体基因（matK和rbcL）和一个核糖体基因（18S rDNA）的序列分析,对代表了基部被子植物和单子叶植物主要谱系分支的86科126属151种被子植物（单子叶植物58科86属101种）进行了系统演化关系分析。研究结果表明由胡椒目Piperales、樟目Laurales、木兰目Magnoliales和林仙目Canellales构成的真木兰类复合群是单子叶植物的姐妹群。单子叶植物的单系性在3个序列联合分析中得到98％的强烈自展支持。联合分析鉴定出9个单子叶植物主要谱系（广义泽泻目Alismatales、薯蓣目Dioscorcales、露兜树目Pandanales、天门冬目Asparagalcs、百合目Liliales、棕榈目Arecales、禾本目Poales、姜目Zingiberales、鸭跖草目Commelinales）和6个其他被子植物主要谱系（睡莲目Nymphaeales、真双子叶植物、木兰目、樟目、胡椒目、林仙目）。在单子叶植物内,菖蒲目Acorales（菖蒲属Acorus）是单子叶植物最早分化的一个谱系,广义泽泻目（包括天南星科Araceae和岩菖蒲科Toficldiaccae）紧随其后分化出来,二者依次和其余单子叶植物类群构成姐妹群关系。无叶莲科Petrosaviaceac紧随广义的泽泻目之后分化出来,无叶莲科和剩余的单子叶植物类群形成姐妹群关系,并得到了较高的支持率。继无叶莲科之后分化的类群形成两个大的分支：一支是由露兜树目和薯蓣目构成,二者形成姐妹群关系：另一支是由天门冬目、百合目和鸭跖草类复合群组成,三者之间的关系在单个序列分析和联合分析中不稳定,需要进一步扩大取样范围来确定。在鸭跖草类复合群分支内,鸭跖草目和姜目的姐妹群关系在3个序列联合分析和2个序列联合分析的严格一致树中均得到强烈的自展支持,获得的支持率均是100％。但是,对于棕榈目和禾本目在鸭跖草类中的系统位置以及它们和鸭跖草目-姜目之间的关系,有待进一步解决。值得注意的是,无叶莲科与其他单子叶植物类群（除菖蒲目和泽泻目外）的系统关系在本文中获得较高的自展支持率,薯蓣目和天门冬目的单系性在序列联合分析中都得到了较好的自展支持,而这些在以往的研究中通常支持率较低。鉴于菖蒲科和无叶莲科独特的系统演化位置,本文支持将其分别独立成菖蒲目和无叶莲目Petrosavialcs的分类学界定。     

Monocot relationships: an overview

In 10 years, the monocots have gone from being one of the least studied and most phylogenetically misunderstood groups of the angiosperms to one of the best characterized. Based on analyses of seven genes representing all three genomes, the following clades have high bootstrap support: Acorales (with the single genus Acorus) is sister to the rest of the monocots, followed successively by Alismatales (including Araceae and Tofieldiaceae), Petrosaviales, Dioscoreales/Pandanales, Liliales, Asparagales, and finally a polytomy of Arecales, Commelinales/Zingiberales, Dasypogonaceae, and Poales. Many of these results also have support from at least some morphological data, but some are unique to the trees created from DNA sequence data. Monocots have been shown in molecular clock studies to be at least 140 million years old, and all major clades and most families date to well before the end of the Cretaceous. More data are required to clarify the positions of the remaining unclearly placed orders, Asparagles, Liliales, and Arecales, as well as Dasypogonaceae. More sequences from the nuclear and mitochondrial genomes are also needed to complement those from the plastid genome, which is the most sampled and thus far most pattern-rich.     

Floral development of petaloid Alismatales as an insight into the origin of the trimerous Bauplan in monocot flowers

Monocots are remarkably homogeneous in sharing a common trimerous pentacyclic floral Bauplan. A major factor affecting monocot evolution is the unique origin of the clade from basal angiosperms. The origin of the floral Bauplan of monocots remains controversial, as no immediate sister groups with similar structure can be identified among basal angiosperms, and there are several possibilities for an ancestral floral structure, including more complex flowers with higher stamen and carpel numbers, or strongly reduced flowers. Additionally, a stable Bauplan is only established beyond the divergence of Alismatales. Here, we observed the floral development of five members of the three ‘petaloid’ Alismatales families Butomaceae, Hydrocharitaceae, and Alismataceae. Outer stamen pairs can be recognized in mature flowers of Alismataceae and Butomaceae. Paired stamens always arise independently, and are either shifted opposite the sepals or close to the petals. The position of stamen pairs is related to the early development of the petals. In Butomaceae, the perianth is not differentiated and the development of the inner tepals is not delayed; the larger inner tepals (petals) only permit the initiation of stamens in antesepalous pairs. Alismataceae has delayed petals and the stamens are shifted close to the petals, leading to an association of stamen pairs with petals in so-called stamen–petal complexes. In the studied Hydrocharitaceae species, which have the monocot floral Bauplan, paired stamens are replaced by larger single stamens and the petals are not delayed. These results indicate that the origin of the floral Bauplan, at least in petaloid Alismatales, is closely linked to the position of stamen pairs and the rate of petal development. Although the petaloid Alismatales are not immediately at the base of monocot divergence, the floral evolution inferred from the results should be a key to elucidate the origin of the floral Bauplan of monocots.     

Complete chloroplast genome sequence of Elodea canadensis and comparative analyses with other monocot plastid genomes

Elodea canadensis is an aquatic angiosperm native to North America. It has attracted great attention due to its invasive nature when transported to new areas in its non-native range. We have determined the complete nucleotide sequence of the chloroplast (cp) genome of Elodea. Taxonomically Elodea is a basal monocot, and only few monocot cp genomes representing early lineages of monocots have been sequenced so far. The genome is a circular double-stranded DNA molecule 156,700bp in length, and has a typical structure with large (LSC 86,194bp) and small (SSC 17,810bp) single-copy regions separated by a pair of inverted repeats (IRs 26,348bp each). The Elodea cp genome contains 113 unique genes and 16 duplicated genes in the IR regions. A comparative analysis showed that the gene order and organization of the Elodea cp genome is almost identical to that of Amborella trichopoda, a basal angiosperm. The structure of IRs in Elodea is unique among monocot species with the whole cp genome sequenced. In Elodea and another monocot Lemna minor the borders between IRs and LSC are located upstream of rps19 gene and downstream of trnH-GUG gene, while in most monocots, IR has extended to include both trnH and rps19 genes. A phylogenetic analysis conducted using Bayesian method, based on the DNA sequences of 81 chloroplast genes from 17 monocot taxa provided support for the placement of Elodea together with Lemna as a basal monocot and the next diverging lineage of monocots after Acorales. In comparison with other monocots, the Elodea cp genome has gone through only few rearrangements or gene losses. IR of Elodea has a unique structure among the monocot species studied so far as its structure is similar to that of a basal angiosperm Amborella. This result together with phylogenetic analyses supports the placement of Elodea as a basal monocot to the next diverging lineage of monocots after Acorales. So far, only few cp genomes representing early lineages of monocots have been sequenced and, therefore, this study provides valuable information about the course of evolution in divergence of monocot lineages.     

Embryology of <Emphasis Type="Italic">Petrosavia</Emphasis> (Petrosaviaceae,Petrosaviales): evidence for the distinctness of the family from other monocots

The affinities of Petrosavia, a rare, leafless, mycoheterotrophic genus composed of two species indigenous to East to Southeast Asia, have long been uncertain. However, recent molecular analyses show that the genus is sister to Japonolirion osense. Japonolirion and Petrosavia comprise the Petrosaviaceae, which are now placed in its own order, Petrosaviales, distinct from other monocots based on molecular analyses. We conducted an embryological study of Petrosavia, comparing it to Japonolirion, as well as to basal monocots (Acorus and Araceae) and more derived monocots (Nartheciaceae, Velloziaceae, and Triuridaceae). Our results showed that Petrosavia is very similar in embryology to Japonolirion, with both genera sharing a glandular anther tapetum, simultaneous cytokinesis in microspore mother cells, anatropous and crassinucellate ovules, T-shaped tetrads of megaspores, ab initio Cellular-type endosperm, and a mature seed coat composed of the exotesta, endotesta, and endotegmen. The two genera of Petrosaviaceae are clearly distinct from Acorus, and all Araceae, Nartheciaceae, Velloziaceae, and Triuridaceae genera in various combinations of characters. Thus, both molecular and embryological evidence support the distinctness of the Petrosaviaceae from other monocots and its placement in its own order, Petrosaviales.     

Systematic embryology of the Araceae

Embryological data of systematic significance to the family Araceae are reviewed and analyzed, with special attention given to the determination of character-state polarities. Character-states considered primitive within the family include: presence of endothecial thickenings; binucleate, starchless pollen; anatropous, crassinucellate ovules with a thick nucellar cap and a single, unbranched funicular bundle; solanad or caryophyllad embryogeny; helobial endosperm development, with 2–8 cells in the chalazal chamber andab initio cellular division in the micropylar chamber; and endosperm present in ripe seeds. The phylogenetic implications of these conclusions are discussed, and promising avenues for future research are indicated.     

Molecular phylogeny of monocotyledons inferred from combined analysis of plastid<Emphasis Type="Italic"> matK</Emphasis> and<Emphasis Type="Italic"> rbcL</Emphasis> gene sequences

Using matK and rbcL sequences (3,269 bp in total) from 113 genera of 45 families, we conducted a combined analysis to contribute to the understanding of major evolutionary relationships in the monocotyledons. Trees resulting from the parsimony analysis are similar to those generated by earlier single or multiple gene analyses, but their strict consensus tree provides much better resolution of relationships among major clades. We find that Acorus (Acorales) is a sister group to the rest of the monocots, which receives 100% bootstrap support. A clade comprising Alismatales is diverged as the next branch, followed successively by Petrosaviaceae, the Dioscoreales–Pandanales clade, Liliales, Asparagales and commelinoids. All of these clades are strongly supported (with more than 90% bootstrap support). The sister-group relationship is also strongly supported between Alismatales and the remaining monocots (except for Acorus) (100%), between Petrosaviaceae and the remaining monocots (except for Acorus and Alismatales) (100%), between the clade comprising Dioscoreales and Pandanales and the clade comprising Liliales, Asparagales and commelinoids (87%), and between Liliales and the Asparagales–commelinoids clade (89%). Only the sister-group relationship between Asparagales and commelinoids is weakly supported (68%). Results also support the inclusion of Petrosaviaceae in its own order Petrosaviales, Nartheciaceae in Dioscoreales and Hanguanaceae in Commelinales.Electronic Supplementary Material Supplementary material is available in the online version of this article at http://dx.doi.org/10.1007/s10265-003-0133-3     

Phylogenetic tree of vascular plants reveals the origins of aquatic angiosperms

Although aquatic plants are discussed as a unified biological group, they are phylogenetically well dispersed across the angiosperms. In this study, we annotated the aquatic taxa on the tree of vascular plants, and extracted the topology of these aquatic lineages to construct the tree of aquatic angiosperms. We also reconstructed the ancestral areas of aquatic families. We found that aquatic angiosperms could be divided into two different categories: the four aquatic orders and the aquatic taxa in terrestrial orders. Aquatic lineages evolved early in the radiation of angiosperms, both in the orders Nymphaeales and Ceratophyllales and among basal monocots (Acorales and Alismatales). These aquatic orders do not have any extant terrestrial relatives. They originated from aquatic habitats during the Early Cretaceous. Asia would have been one of the centers for early diversification of aquatic angiosperms. The aquatic families within terrestrial orders may originate from other areas besides Asia, such as America or Australia. The lineages leading to extant angiosperms diversified early in underexploited freshwater habitats. The four extant aquatic orders were relicts of an early radiation of angiosperm in aquatic environments. Their extinct ancestors might be aquatic early angiosperms.     

EVOLUTIONARY AND ECOLOGICAL SIGNIFICANCE OF STARCH STORAGE IN POLLEN OF THE ARACEAE

Starch content was qualitatively assessed for pollen of 79 of the 111 currently recognized genera of the family Araceae—one of three monocot families known to exhibit both starchy and starchless pollen. Although 73% of the genera investigated had exclusively starchy pollen, character correlation suggests that starchless pollen is the primitive type for the family Araceae, as well as for monocots in general. Pollen starch content is a highly conservative character at the generic level in Araceae; only a single genus (Schismatoglottis) clearly exhibits both character states. The distribution of starchy pollen among aroid genera is consistent with what have here been termed Bakers' Starch Laws. Aroid pollen below a certain critical diameter—17-25 μm—is almost invariably starchless. Larger pollen is nearly always starchy, except where insect pollinators may use pollen nutritionally. There is strong evidence that the trend from starchless to starchy pollen in Araceae is reversible, according to the constraints imposed by the aforementioned factors.     

Evolutionary timescale of monocots determined by the fossilized birth‐death model using a large number of fossil records

Although the phylogenetic relationships between monocot orders are sufficiently understood, a timescale of their evolution is needed. Several studies on molecular clock dating are available, but their results have been biased by their calibration schemes. Recently, the fossilized birth‐death model, a type of Bayesian dating method, was proposed, and it does not require prior calibration and allows the use all available fossils. Using this model, we conducted divergence‐time estimations of monocots to explore their evolutionary timeline without calibration bias. This is the first application of this model to seed plants. The dataset contained the matK and rbcL chloroplast genes of 118 monocot genera covering all extant orders. We employed information from 247 monocot fossils, which exceeded previous dating analyses that used a maximum of 12 monocot fossils. The crown group of monocots was dated to approximately the Late Jurassic–Early Cretaceous periods, and most extant monocot orders were estimated to diverge throughout the Early Cretaceous. Our results overlapped with the divergence time of insect lineages, such as beetles and flies, suggesting an association with pollinators in early monocot evolution. In addition, we proposed three new orders based on divergence time: Orchidales separated from Asparagales and Tofieldiales and Arales separated from Aslimatales.     

Seed fertilization, development, and germination in Hydatellaceae (Nymphaeales): Implications for endosperm evolution in early angiosperms

New data on endosperm development in the early-divergent angiosperm Trithuria (Hydatellaceae) indicate that double fertilization results in formation of cellularized micropylar and unicellular chalazal domains with contrasting ontogenetic trajectories, as in waterlilies. The micropylar domain ultimately forms the cellular endosperm in the dispersed seed. The chalazal domain forms a single-celled haustorium with a large nucleus; this haustorium ultimately degenerates to form a space in the dispersed seed, similar to the chalazal endosperm haustorium of waterlilies. The endosperm condition in Trithuria and waterlilies resembles the helobial condition that characterizes some monocots, but contrasts with Amborella and Illicium, in which most of the mature endosperm is formed from the chalazal domain. The precise location of the primary endosperm nucleus governs the relative sizes of the chalazal and micropylar domains, but not their subsequent developmental trajectories. The unusual tissue layer surrounding the bilobed cotyledonary sheath in seedlings of some species of Trithuria is a belt of persistent endosperm, comparable with that of some other early-divergent angiosperms with a well-developed perisperm, such as Saururaceae and Piperaceae. The endosperm of Trithuria is limited in size and storage capacity but relatively persistent.     

The Mitochondrial Genome of an Aquatic Plant,Spirodela polyrhiza

Background

Spirodela polyrhiza is a species of the order Alismatales, which represent the basal lineage of monocots with more ancestral features than the Poales. Its complete sequence of the mitochondrial (mt) genome could provide clues for the understanding of the evolution of mt genomes in plant.

Methods

Spirodela polyrhiza mt genome was sequenced from total genomic DNA without physical separation of chloroplast and nuclear DNA using the SOLiD platform. Using a genome copy number sensitive assembly algorithm, the mt genome was successfully assembled. Gap closure and accuracy was determined with PCR products sequenced with the dideoxy method.

Conclusions

This is the most compact monocot mitochondrial genome with 228,493 bp. A total of 57 genes encode 35 known proteins, 3 ribosomal RNAs, and 19 tRNAs that recognize 15 amino acids. There are about 600 RNA editing sites predicted and three lineage specific protein-coding-gene losses. The mitochondrial genes, pseudogenes, and other hypothetical genes (ORFs) cover 71,783 bp (31.0%) of the genome. Imported plastid DNA accounts for an additional 9,295 bp (4.1%) of the mitochondrial DNA. Absence of transposable element sequences suggests that very few nuclear sequences have migrated into Spirodela mtDNA. Phylogenetic analysis of conserved protein-coding genes suggests that Spirodela shares the common ancestor with other monocots, but there is no obvious synteny between Spirodela and rice mtDNAs. After eliminating genes, introns, ORFs, and plastid-derived DNA, nearly four-fifths of the Spirodela mitochondrial genome is of unknown origin and function. Although it contains a similar chloroplast DNA content and range of RNA editing as other monocots, it is void of nuclear insertions, active gene loss, and comprises large regions of sequences of unknown origin in non-coding regions. Moreover, the lack of synteny with known mitochondrial genomic sequences shed new light on the early evolution of monocot mitochondrial genomes.     

Microsporogenesis in Monocotyledons

This paper critically reviews the distribution of microsporogenesistypes in relation to recent concepts in monocot systematics.Two basic types of microsporogenesis are generally recognized:successive and simultaneous, although intermediates occur. Theseare characterized by differences in tetrad morphology, generallytetragonal or tetrahedral, although other forms occur, particularlyassociated with successive division. Successive microsporogenesisis predominant in monocotyledons, although the simultaneoustype characterizes the ‘lower’ Asparagales. Simultaneousmicrosporogenesis also occurs inJaponolirion and Petrosavia(unplaced taxa), some Araceae, Aponogeton, Thalassia andTofieldia(Alismatales), Dioscorea, Stenomeris and Tacca (Dioscoreales),and some Commelinanae: Arecaceae (Arecales), and Cyperaceae,Juncaceae and Thurniaceae (Poales). Simultaneous microsporogenesisis of phylogenetic significance within some of these groups,for example, Asparagales, Dioscoreales and Poales. An intermediatetype is recorded in Stemonaceae (Pandanales), Commelinaceae(Commelinales) and in Eriocaulaceae and Flagellariaceae (Poales).There is little direct relationship between microsporogenesistype and pollen aperture type in monocots (except for trichotomosulcateand pantoporate apertures), although trichotomosulcate aperturesin monocot pollen, and equatorial tricolpate and tricolporateapertures in eudicot pollen, are all related to simultaneousmicrosporogenesis. Copyright 1999 Annals of Botany Company Microsporogenesis, monocotyledons, pollen apertures, phylogeny, tetrads, simultaneous, successive, systematics.     

Dynamics and evolution of the inverted repeat-large single copy junctions in the chloroplast genomes of monocots

Background  

Various expansions or contractions of inverted repeats (IRs) in chloroplast genomes led to fluxes in the IR-LSC (large single copy) junctions. Previous studies revealed that some monocot IRs contain a trnH-rps19 gene cluster, and it has been speculated that this may be an evidence of a duplication event prior to the divergence of monocot lineages. Therefore, we compared the organizations of genes flanking two IR-LSC junctions in 123 angiosperm representatives to uncover the evolutionary dynamics of IR-LSC junctions in basal angiosperms and monocots.