释百合科(广义)分类的沿革及对我国今后研究的展望

本文试图阐明近60年来一些作者对广义百合科所作的重大变革。广义百合科分类的变革不能不涉及其近缘科的分类,因此我们以Engler＆Prantl(1930)的Liliiflorae目中的族作为基本单位,列表以明Hutchinson(1934)、Dahlgren et al．(1985,包括G．Dahlgren,989)、Takhtajan(1987)和Thorne(1992)各系统所作不同的处理。作者对Hutchinson的观点和其采用的分类特征作了较详细的介绍,因为他可能是对广义百合科作重大变革的第一个作者,而对Dahlgen et al.、Takhtajan和Thorne系统的介绍则较为简略。本文还讨论了科级范围大小的利弊,不同意有些作者仅仅将亚科或卫星类群都提升为科的作法。假如这种提升之后仍如原先的科保留于同一类群之中,而不作系统上的改变,则意义不大。 本文对Hutchinson、Takhtajan和Dahlgren et al．(1985)的单子叶植物起源的假说也作了介绍。虽然Dahlgren et al．的见解未被大多数作者所接受,但他们的方法论值得注意。基于单子叶植物和双子叶植物的共性,他们假设出一个臆想的分类群,从这个臆想的分类群演进到一个单子叶植物祖先只需经过最少的步段。这样的一个单子叶植物祖先现在是不存在的,但最接近它的是现存的Trichopus(隶Trichopodaceae,Dioscordales)。Trichopus肯定不是单子叶植物和双子叶植物之间的一个直接联系的纽带,但它有许多特征可以在Annonales(隶lianae超目)中找到。因此,他们认为Dioscoreales是最不特化的单子叶植物,把它作为百合超目的第一个目,置于单子叶植物系统的最前位置。本文作者最后对今后我国开展广义百合科研究提出一些建议,其中以选择一些分布于北半球类群,如Lilium,Monocharis,Paris等等,作广泛研究(可从形态学、细胞学、分子系统学三个层次进行)最为现实,因为对它们取材比较容易。     

Young,intact and nested retrotransposons are abundant in the onion and asparagus genomes

Background and Aims

Although monocotyledonous plants comprise one of the two major groups of angiosperms and include >65 000 species, comprehensive genome analysis has been focused mainly on the Poaceae (grass) family. Due to this bias, most of the conclusions that have been drawn for monocot genome evolution are based on grasses. It is not known whether these conclusions apply to many other monocots.

Methods

To extend our understanding of genome evolution in the monocots, Asparagales genomic sequence data were acquired and the structural properties of asparagus and onion genomes were analysed. Specifically, several available onion and asparagus bacterial artificial chromosomes (BACs) with contig sizes >35 kb were annotated and analysed, with a particular focus on the characterization of long terminal repeat (LTR) retrotransposons.

Key Results

The results reveal that LTR retrotransposons are the major components of the onion and garden asparagus genomes. These elements are mostly intact (i.e. with two LTRs), have mainly inserted within the past 6 million years and are piled up into nested structures. Analysis of shotgun genomic sequence data and the observation of two copies for some transposable elements (TEs) in annotated BACs indicates that some families have become particularly abundant, as high as 4–5 % (asparagus) or 3–4 % (onion) of the genome for the most abundant families, as also seen in large grass genomes such as wheat and maize.

Conclusions

Although previous annotations of contiguous genomic sequences have suggested that LTR retrotransposons were highly fragmented in these two Asparagales genomes, the results presented here show that this was largely due to the methodology used. In contrast, this current work indicates an ensemble of genomic features similar to those observed in the Poaceae.     

Asparagales Telomerases which Synthesize the Human Type of Telomeres

The order of monocotyledonous plants Asparagales is attractive for studies of telomere evolution as it includes three phylogenetically distinct groups with telomeres composed of TTTAGGG (Arabidopsis-type), TTAGGG (human-type) and unknown alternative sequences, respectively. To analyze the molecular causes of these switches in telomere sequence (synthesis), genes coding for the catalytic telomerase subunit (TERT) of representative species in the first two groups have been cloned. Multiple alignments of the sequences, together with other TERT sequences in databases, suggested candidate amino acid substitutions grouped in the Asparagales TERT synthesizing the human-type repeat that could have contributed to the changed telomere sequence. Among these, mutations in the C motif are of special interest due to its functional importance in TERT. Furthermore, two different modes of initial elongation of the substrate primer were observed in Asparagales telomerases producing human-like repeats, which could be attributed to interactions between the telomerase RNA subunit (TR) and the substrate. Electronic Supplementary Material Supplementary material is available for this article at and is accessible for authorized users.     

The historical evolutionary development of Hemerocallis middendorfii (Hemerocallidaceae) revealed by non-coding regions in chloroplast DNA

A procedure in which combined molecular phylogeographical analyses among populations of Hemerocallis middendorfii (Hemerocallidaceae, Asparagales) were applied allowed comparisons to be made with the geological history. Information on geographical areas in which synapomorphic mutations occurred was used in molecular phylogeographical analyses for the first time in this study, in addition to molecular maximum-parsimonious analyses and TCS analyses. Nucleotide sequences of the intergenic region between the rbcL and atpB genes and the trnL (UAA) intron in chloroplast DNA were analyzed for 28 Japanese and three Chinese populations of H. middendorfii, 10 populations of closely related species and four outgroup species. The data of the three analyses in general agreed with one another and indicated that the separation of this species had proceeded in conjunction with the geological vicariance or orogeny. Where the synapomorphic mutations occurred, three patterns were recognized. They correlated with the geological history, namely, the age when the Japanese Archipelago and the Continent had joined, when the Sea of Japan was formed, and when there was separation from the Continent and the formation of each of the islands in the Japanese Archipelago. It was suggested that ancestors of H. middendorfii originated at latest 25 million years ago when the Japanese Archipelago and the Continent had joined; that is, before the formation of the Sea of Japan.We are greatly obliged to three anonymous reviewers for their constructive comments.     

Phylogenetic Relationships of Amaryllidaceae Based on matK Sequence Data

matK gene, which is located in the chloroplast genome and evolves more quickly than the rbcL gene. A total of 31 species representing 31 of the 59 genera in the family were examined in this study. We also used 21 species from another ten families of Asparagales, four species from three families of Liliales and Acorus as outgroups. We obtained partial sequences of matK with lengths of 1,109–1,148 bp, corresponding to positions 230 to 1,343 of the Oryza sativa matK gene. The pairwise percentage sequence divergence ranged from 0 to 19.1% for all the species examined except Acorus, and 0 to 4.6% within Amaryllidaceae. Two methods of phylogenetic analysis, the Maximum Parsimony and Neighbor-Joining methods, were used. The trees obtained from these two analyses were fundamentally consistent. In both trees, the Amaryllidaceae sensu Dahlgren et al. formed a well-supported monophyletic clade with 100% bootstrap support. Amaryllidaceae were included in the Asparagales; however, its phylogenetic position within the Asparagales was not clearly resolved. Judging from the NJ tree, Agapanthus might be a sister group of the Amaryllidaceae, although bootstrap support for this was low. Character-state mapping was used to infer a center of origin and the biogeographic history of Amaryllidaceae. The result supports the hypothesis that the family evolved in Africa and subsequently spread to other continents, further suggesting that South America is the center of secondary diversification. Received 6 January 1999/ Accepted in revised form 8 April 1999     

Floral structure and development and systematic aspects of some 'lower' Asparagales

 Floral structure and development of representatives of Asteliaceae, Blandfordiaceae, Boryaceae, Doryanthaceae, and Hypoxidaceae, all members of the `lower' Asparagales, were studied comparatively. The results are discussed in the light of new molecular systematic studies, but also with regard to established morphological characters in related groups. Stamen shape varies considerably within and between taxa: the shape of anthers is from X-shaped, sagittate to non-sagittate, they are either latrorse or introrse, basifixed, centrifixed or dorsifixed. Gynoecia are syncarpous up to the stigmatic region in all taxa. Ovaries of Doryanthaceae and Hypoxidaceae are inferior, but they are superior in Asteliaceae, Blandfordiaceae and Boryaceae. All ovaries have at least a short synascidiate zone. With the exception of Astelia alpina (Asteliaceae), the ovaries are trilocular. Ovaries of Asteliaceae contain mucilage, which is secreted from trichomes on the funicle and on the placenta. Although flowers are polysymmetric at anthesis, they are monosymmetric in earliest stages with a developmental gradient from adaxial to abaxial. Perianth organs arise individually from either a concave (taxa with inferior ovary) or convex (taxa with superior ovary) apex. Hypoxidaceae have pollen flowers with free stamens. One species, Curculigo capitulata, has Solanum-type flowers with postgenitally united stamens. It is most probably pollinated by buzzing bees. All other taxa have nectariferous flowers with internal or external septal nectaries. Received February 5, 2001 Accepted June 20, 2001     

Anatomical and molecular systematics of Asteliaceae and Hypoxidaceae

The astelioid group of asparagoid lilies (Lilianae - Asparagales) comprises Hypoxidaceae, Asteliaceae, Blandfordia and Lanaria. New information is presented on astelioid anatomy, together with a review of other systematic characters. These data are analysed in the context of recent evidence from rbc L nucleotide sequences that astelioids are related to orchids, and that astelioids and orchids (plus Alania and Borya ) form a clade that is sister to all other asparagoid taxa. Hypoxidaceae and Asteliaceae differ from each other in several respects, but there are certain characters linking the two families, notably branched hairs and mucilage canals, unusual characters in Lilianae. Family diagnoses are upheld, but the precise relationships of Blandfordia and Lanaria are still poorly supported within the astelioid clade.     

Multiple developmental pathways leading to a single morph: monosulcate pollen (examples from the Asparagales)

BACKGROUND AND AIMS: Early developmental events in microsporogenesis are known to play a role in pollen morphology: variation in cytokinesis type, cell wall formation, tetrad shape and aperture polarity are responsible for pollen aperture patterning. Despite the existence of other morphologies, monosulcate pollen is one of the most common aperture types in monocots, and is also considered as the ancestral condition in this group. It is known to occur from either a successive or a simultaneous cytokinesis. In the present study, the developmental sequence of microsporogenesis is investigated in several species of Asparagales that produce such monosulcate pollen, representing most families of this important monocot clade. METHODS: The developmental pathway of microsporogenesis was investigated using light transmission and epifluorescence microscopy for all species studied. Confocal microscopy was used to confirm centripetal cell plate formation. KEY RESULTS: Microsporogenesis is diverse in Asparagales, and most variation is generally found between families. It is confirmed that the whole higher Asparagales clade has a very conserved microsporogenesis, with a successive cytokinesis and centrifugal cell plate formation. Centripetal cell wall formation is described in Tecophilaeaceae and Iridaceae, a feature that had so far only been reported for eudicots. CONCLUSIONS: Monosulcate pollen can be obtained from several developmental pathways, leading thus to homoplasy in the monosulcate character state. Monosulcate pollen should not therefore be considered as the ancestral state unless it is produced through the ancestral developmental pathway. The question about the ancestral developmental pathway leading to monosulcy remains open.     

The discovery of polyandry in Curculigo (Hypoxidaceae): implications for androecium evolution of asparagoid monocotyledons

BACKGROUND AND AIMS: Individual flowers of the monocot Curculigo racemosa (Hypoxidaceae, Asparagales) are regularly polyandrous. To evaluate the significance of this almost unique character among Asparagales for flower evolution of asparagoid monocots, flowers of C. racemosa were studied comparatively. METHODS: Anthetic flowers as well as early floral developmental stages were studied by light and scanning electron microscopy. KEY RESULTS: Despite the polyandry, floral development is similar to that of other Asparagales with a developmental gradient from adaxial to abaxial. Stamens initiate simultaneously and the diameter of staminal primordia is about half of that in species with six anthers. The number of stamens is not fixed (12-26) and varies within the same inflorescence. Surprisingly, the gynoecium can be four- or six-locular, besides the normal trimerous state. CONCLUSIONS: The discovery of a polyandrous Curculigo reveals plasticity of stamen number at the base of Asparagales. Orchidaceae - sister to all other Asparagales - has a reduced stamen number (three, two or one), whereas in Hypoxidaceae - part of the next diverging clade - either the normal monocot stamen number (six), polyandry (this study) or the loss of three anthers (Pauridia) occurs. However, at present it is impossible to decide whether the flexibility in stamen number is autapomorphic for each group or whether it is a synapomorphy. The small size of stamen primordia of Curculigo is conspicuous. It allows more space for additional androecial primordia. Stamens are initiated as independent organs, and filaments are not in bundles, hence C. racemosa is not secondarily polyandrous as may be the case in the distantly related Gethyllis of asparagoid Amaryllidaceae. The increase in carpel number is a rare phenomenon in angiosperms. A possible explanation for the polyandry of C. racemosa is that it is a natural SUPERMAN-deficient mutant, which shows an increase of stamens, or ULTRAPETALA or CARPEL FACTORY mutants, which are polyandrous and changed in carpel number.