Phytohormones and Cytoskeleton

Various aspects of the interaction between phytohormones and principal components of the cytoskeleton are considered. Four levels of this interaction are discussed: (1) the spatial organization of microtubules and microfilaments; (2) the synthesis and modification of cytoskeletal proteins; (3) the cytoskeleton involvement in processes, which are under hormonal control; and (4) the cytoskeleton involvement in phytohormone functioning.     

Alkylation damage in DNA and RNA--repair mechanisms and medical significance

Alkylation lesions in DNA and RNA result from endogenous compounds, environmental agents and alkylating drugs. Simple methylating agents, e.g. methylnitrosourea, tobacco-specific nitrosamines and drugs like temozolomide or streptozotocin, form adducts at N- and O-atoms in DNA bases. These lesions are mainly repaired by direct base repair, base excision repair, and to some extent by nucleotide excision repair (NER). The identified carcinogenicity of O(6)-methylguanine (O(6)-meG) is largely caused by its miscoding properties. Mutations from this lesion are prevented by O(6)-alkylG-DNA alkyltransferase (MGMT or AGT) that repairs the base in one step. However, the genotoxicity and cytotoxicity of O(6)-meG is mainly due to recognition of O(6)-meG/T (or C) mispairs by the mismatch repair system (MMR) and induction of futile repair cycles, eventually resulting in cytotoxic double-strand breaks. Therefore, inactivation of the MMR system in an AGT-defective background causes resistance to the killing effects of O(6)-alkylating agents, but not to the mutagenic effect. Bifunctional alkylating agents, such as chlorambucil or carmustine (BCNU), are commonly used anti-cancer drugs. DNA lesions caused by these agents are complex and require complex repair mechanisms. Thus, primary chloroethyl adducts at O(6)-G are repaired by AGT, while the secondary highly cytotoxic interstrand cross-links (ICLs) require nucleotide excision repair factors (e.g. XPF-ERCC1) for incision and homologous recombination to complete repair. Recently, Escherichia coli protein AlkB and human homologues were shown to be oxidative demethylases that repair cytotoxic 1-methyladenine (1-meA) and 3-methylcytosine (3-meC) residues. Numerous AlkB homologues are found in viruses, bacteria and eukaryotes, including eight human homologues (hABH1-8). These have distinct locations in subcellular compartments and their functions are only starting to become understood. Surprisingly, AlkB and hABH3 also repair RNA. An evaluation of the biological effects of environmental mutagens, as well as understanding the mechanism of action and resistance to alkylating drugs require a detailed understanding of DNA repair processes.     

13C-, 15N- and 31P-NMR studies of oxidized and reduced low molecular mass thioredoxin reductase and some mutant proteins.

Thioredoxin reductase (TrxR) from Escherichia coli, the mutant proteins E159Y and C138S, and the mutant protein C138S treated with phenylmercuric acetate were reconstituted with [U-(13)C(17),U-(15)N(4)]FAD and analysed, in their oxidized and reduced states, by (13)C-, (15)N- and (31)P-NMR spectroscopy. The enzymes studied showed very similar (31)P-NMR spectra in the oxidized state, consisting of two peaks at -9.8 and -11.5 p.p.m. In the reduced state, the two peaks merge into one apparent peak (at -9.8 p.p.m.). The data are compared with published (31)P-NMR data of enzymes closely related to TrxR. (13)C and (15)N-NMR chemical shifts of TrxR and the mutant proteins in the oxidized state provided information about the electronic structure of the protein-bound cofactor and its interactions with the apoproteins. Strong hydrogen bonds exist between protein-bound flavin and the apoproteins at C(2)O, C(4)O, N(1) and N(5). The N(10) atoms in the enzymes are slightly out of the molecular plane of the flavin. Of the ribityl carbon atoms C(10alpha,gamma,delta) are the most affected upon binding to the apoprotein and the large downfield shift of the C(10gamma) atom indicates strong hydrogen bonding with the apoprotein. The hydrogen bonding pattern observed is in excellent agreement with X-ray data, except for the N(1) and the N(3) atoms where a reversed situation was observed. Some chemical shifts observed in C138S deviate considerably from those of the other enzymes. From this it is concluded that C138S is in the FO conformation and the others are in the FR conformation, supporting published data. In the reduced state, strong hydrogen bonding interactions are observed between C(2)O and C(4)O and the apoprotein. As revealed by the (15)N chemical shifts and the N(5)H coupling constant the N(5) and the N(10) atom are highly sp(3) hybridized. The calculation of the endocyclic angles for the N(5) and the N(10) atoms shows the angles to be approximately 109 degrees, in perfect agreement with X-ray data showing that the flavin assumes a bent conformation along the N(10)/N(5) axis of the flavin. In contrast, the N(1) is highly sp(2) hybridized and is protonated, i.e. in the neutral state. Upon reduction of the enzymes, the (13)C chemical shifts of some atoms of the ribityl side chain undergo considerable changes also indicating conformational rearrangements of the side-chain interactions with the apoproteins. The chemical shifts between native TrxR and C138S are now rather similar and differ from those of the two other mutant proteins. This strongly indicates that the former enzymes are in the FO conformation and the other two are in the FR conformation. The data are discussed briefly in the context of published NMR data obtained with a variety of flavoproteins.     

Water and ion binding around r(UpA)12 and d(TpA)12 oligomers--comparison with RNA and DNA (CpG)12 duplexes

The structural and dynamic properties of the water and ion first coordination shell of the r(A-U) and d(A-T) base-pairs embedded within the r(UpA)12 and d(TpA)12 duplexes are described on the basis of two 2.4 ns molecular dynamics simulations performed in a neutralizing aqueous environment with 0.25 M added KCl. The results are compared to previous molecular dynamics simulations of the r(CpG)12 and d(CpG)12 structures performed under similar conditions. It can be concluded that: (i) RNA helices are more rigid than DNA helices of identical sequence, as reflected by the fact that RNA duplexes keep their initial A-form shape while DNA duplexes adopt more sequence-specific shapes. (ii) Around these base-pairs, the water molecules occupy 21 to 22 well-defined hydration sites, some of which are partially occupied by potassium ions. (iii) These hydration sites are occupied by an average of 21.9, 21.0, 20.1, and 19.8 solvent molecules (water and ions) around the r(G=C), r(A-U), d(G=C), and d(A-T) pairs, respectively. (iv) From a dynamic point of view, the stability of the hydration shell is the strongest for the r(G=C) pairs and the weakest for the d(A-T) pairs. (v) For RNA, the observed long-lived hydration patterns are essentially non-sequence dependent and involve water bridges located in the deep groove and linking OR atoms of adjacent phosphate groups. Maximum lifetimes are close to 400 ps. (vi) In contrast, for DNA, long-lived hydration patterns are sequence dependent and located in the minor groove. For d(CpG)12, water bridges linking the (G)N3 and (C)O2 with the O4' atoms of adjacent nucleotides with 400 ps maximum lifetimes are characterized while no such bridges are observed for d(TpA)12. (vii) Potassium ions are observed to bind preferentially to deep/major groove atoms at RpY steps, essentially d(GpC), r(GpC), and r(ApU), by forming ion-bridges between electronegative atoms of adjacent base-pairs. On average, about half an ion is observed per base-pair. Positive ion-binding determinants are related to the proximity of two or more electronegative atoms. Negative binding determinants are associated with the electrostatic and steric hindrance due to the proximity of electropositive amino groups and neutral methyl groups. Potassium ions form only transient contacts with phosphate groups.     

观赏蕨类引种栽培及其物候期的观察

对中华水韭、福建观音座莲、华南紫萁、西南凤尾蕨、剑叶铁角蕨、翅轴蹄盖蕨、东方荚果蕨、同形鳞毛蕨、圆顶耳蕨等30种蕨类进行了引种栽培和物候期观察.结果表明:(1)引种成活率高;(2)管理粗放;(3)多数种类四季常青,形态优美,具有较高的观赏价值,能够在怀化市安全越冬,值得在亚热带地区开发利用;(4)华南紫萁、小黑桫椤、光蹄盖蕨、长江蹄盖蕨、翅轴蹄盖蕨、三相蕨、同形鳞毛蕨、圆顶耳蕨、镰羽贯众等9种蕨类在展叶时,同时长出孢子囊;(5)多数蕨类的孢子囊在长出后1个月左右发育成熟;(6)在叶开始萌动时引种栽培的蕨类,当年的营养叶萌发期和展叶期都推迟半个月左右,但孢子囊群的出现期和成熟期不受影响;(7)不同蕨类的孢子囊群形成期具有差异性,同种蕨类的孢子囊群形成期具有相对稳定性,因此,可以作为鉴别物种的依据之一。建议在编写、修订《中国植物志》和地方植物志时增加蕨类孢子囊群形成期的描述。     

The types of homology and their significance for evolutionary biology and phylogenetics

This paper comments on recently revived discussion about the most adequate definition of homology. Homologues are considered as similarities of complex structures or patterns which are based on a continuity of biological information or instruction. Dependent on the level of comparison four types of homology are defined: (1) Iterative ( = serial = homonomy), (2) ontogenetic, (3) di- or polymorphic, and (4) supraspecific homology. The significance of all four types for evolutionary biology and phylogenetic analysis is outlined.     

Rates and Patterns of Scndna and Mtdna Divergence within the Drosophila Melanogaster Subgroup

Levels of DNA divergence among the eight species of the Drosophila melanogaster subgroup and D. takahashii have been determined using the technique of DNA-DNA hybridization. Two types of DNA were used: single-copy nuclear DNA (scnDNA) and mitochondrial DNA (mtDNA). The major findings are: (1) A phylogeny has been derived for the group based on scnDNA which is congruent with chromosomal data, morphology, and behavior. The three homosequential species, simulans, sechellia, and mauritiana, are very closely related; the scnDNA divergence indicate the two island species are a monophyletic group. (2) The rates of change of scnDNA and mtDNA are not greatly different; if anything scnDNA evolves faster than mtDNA. (3) The rates of scnDNA evolution are not closely correlated to chromosomal (inversion) evolution. (4) The Drosophila genome appears to consist of two distinct classes of scnDNA with respect to rate of evolutionary change, a very rapidly evolving fraction and a relatively conservative fraction. (5) The absolute rate of change was estimated to be at least 1.7% nucleotide substitution per one million years. (6) DNA distance estimates based on restriction site variation are correlated with distances based on DNA-DNA hybridization, although the correlation is not very strong.     

Lipoprotein receptors and atherosclerosis

The whole lipoprotein spectrum of human plasma may be divided into atherosclerotic and anti-atherosclerotic lipoproteins. To the first class belong apolipoprotein (apo) B and some apoE-containing lipoproteins of the very-low-density (VLDL), intermediate-density (IDL) and low-density (LDL) lipoprotein fractions. Anti-atherosclerotic lipoproteins are apoA-containing high-density lipoproteins (HDL). Circulating plasma lipoproteins are catabolized mainly by specific cell surface receptors (R) which react with apoB and apoE (B/E-R), for apoE (E-R) or for apoA (HDL-R). Whereas the B/E-R and E-R are responsible for the cellular uptake of lipoproteins and their lipid load by various organs, HDL-R are thought to promote lipid (cholesterol) efflux. There is an additional class of lipoprotein receptors, the so called scavenger-R which are responsible for the removal of altered or degraded lipoproteins for the circulation. Under normal physiological conditions, the concerted action of these receptors warrants efficient lipoprotein turnover and direction into target organs. Derangements of this system, however, may lead to the deposition and accumulation of atherogenic lipids, notably free cholesterol (FC) and cholesteryl esters (CE) in arterial tissue causing atherosclerosis and cardiac death.     

Acacia senegal and Prosopis chilensis-nodulating rhizobia Sinorhizobium arboris HAMBI 2361 and S. kostiense HAMBI 2362 produce tetra- and pentameric LCOs that are N-methylated, O-6-carbamoylated and partially sulfated

Sinorhizobium arboris and S. kostiense are rhizobia that nodulate the tropical leguminous trees Acacia senegal and Prosopis chilensis. The lipochito-oligosaccharidic signalling molecules (LCOs) of S. arboris HAMBI 2361 and S. kostiense HAMBI 2362 were analyzed by mass spectrometry. The major LCOs produced by the strains were shown to be pentameric, acylated with common fatty acids, N-methylated, O-6-carbamoylated and partially sulfated, as are the LCOs characterized to date for other Acacia-nodulating rhizobia. Besides the major LCOs the two strains produced (i) tetrameric LCOs, (ii) LCOs acylated with fatty acids other than those commonly found, (iii) LCOs with only an acyl substituent and (iv) noncarbamoylated LCOs. Production of LCOs (i) to (iii) are novel among Acacia-nodulating rhizobia. The roles of the different structural characteristics of LCOs in the rhizobium-A. senegal symbiosis are discussed. Specific structural features of the LCOs are proposed to be important in the selection of effective nitrogen-fixing rhizobia by A. senegal.     

Phosphinyliminodithiolane insecticides: Novel addition reactions of phosfolan and mephosfolan sulfoxides and sulfones

2-(Diethoxyphosphinylimino)-1,3-dithiolane (phosfolan) and its 4-methyl analog (mephosfolan) are proinsecticides as determined by microsomal mixed-function oxidase (MFO) activation to potent acetylcholinesterase (AChE) inhibitors. They are similarly activated by peracid oxidation which yields the sulfoxide and sulfone derivatives. The hydrolytically unstable S-oxides are irreversible AChE inhibitors that are 160- to 47,000-fold more potent than phosfolan and mephosfolan. MFO S-oxidation is indicated for both proinsecticides by (a) NADPH-dependent increases in potency as AChE inhibitors to an extent expected of sulfoxides, and (b) formation of the S-oxide hydrolysis product diethyl phosphoramidate.     

Biochemical characterisation of mitochondrial and lysosomal particles in foetal and neonatal rats

1. The sedimentation coefficients of the mitochondria are larger in the foetal as compared to the adult rats. For cytochrome oxidase, the values are respectively 18.700 x 10(3) and 13,550 x 10(3) S. 2. Lysosomal sedimentation coefficients are smaller in the foetal as compared to the adults. For beta galactosidase, the values are respectively 5.090 x 10(3) and 10.430 x 10(3) S. 3. It is concluded that the mitochondria are larger and the lysosomes smaller in size in the foetus as compared to the adults. 4. The external membrane of neonatal mitochondria and the membrane of foetal and neonatal lysosomes are more resistant to osmotic swelling.     

Nomenclature of Sarcocystis in the ox and sheep and of fecal Coccidia of the dog and cat.

The nomenclature of Sarcocystis and related protozoan genera is reviewed, and modern diagnoses of the genera Isospora, Toxoplasma, Besnoitia, Sarcocystis, and Frenkelia in the coccidian family Eimeriidae are given. S cruzi (Hasselmann 1926) comb. n., S. hirsuta Moulé 1888, and S. hominis (Railliet and Lucet 1891) comb. n. are recognized in the muscles of the ox Bos taurus; S. ovicanis Heydorn, Gestrich, Melhorn, and Rommel 1975, and S. tenella Railliet 1886 are recognized in the muscles of the sheep Ovis aries; S. bigemina (Stiles 1891) comb. n., S. cruzi, S. ovicanis, S. bertrami Doflein 1901, S. miescheriana (Kühn 1865) Lankester 1882, I. ohioensis Dubey 1975, I. canis Nemeséri 1959, Isospora sp. n. Dubey, and Isospora sp. n. Trayser and Todd are recognized in dog (Canis familiaris) feces; and S. hirsuta, S. tenella, S. muris (Blanchard 1885) Labbé 1899, B. besnoiti (Marotel 1912) Henry 1913, Besnoitia sp. n. Frenkel, T. gondii (Nicolle and Manceaus 1908) Nicolle and Manceaux 1909, T. hammondi (Frenkel 1974) Levine and Nye 1976, I. rivolta (Grassi 1879) Wenyon 1923, and I. felis Wenyon 1923 are recognized in cat (Felis catus) feces. Hoareosporidium Pande, Bhatia, and Chauhan 1972 is considered a synonym of Sarcocystis.     

中国石松类和蕨类植物的多样性与地理分布

物种编目及其科属系统排列是了解生物多样性的基础, 本文采用Flora of China使用的分类系统, 结合最新分子分类学研究成果以及近几年发表的新资料, 对中国石松类和蕨类植物多样性和地理分布数据进行了统计和分析。结果表明中国共有石松类和蕨类植物40科178属2,147种5个亚种118个变种, 其中特有种839个, 占总种数的39.08%。种数最多的5个科依次为鳞毛蕨科(505种, 含种下单位, 下同)、蹄盖蕨科(323种)、水龙骨科(280种)、凤尾蕨科(266种)和金星蕨科(209种); 种数最多的5个属依次为耳蕨属(Polystichum, 209种)、鳞毛蕨属(Dryopteris, 176种)、蹄盖蕨属(Athyrium, 137种)、双盖蕨属(Diplazium, 98种)和凤尾蕨属(Pteris, 97种)。在地理分布上, 种数排名前5的省份为云南(1,365种)、四川(875种)、贵州(838种)、广西(785种)和台湾(779种)。含中国特有石松类和蕨类植物的科属中, 排前3位的科分别为鳞毛蕨科(257种)、蹄盖蕨科(169种)和凤尾蕨科(113种); 排前3位的属为耳蕨属(140种)、蹄盖蕨属(82种)和鳞毛蕨属(61种)。     

Stress and adaptation in conservation genetics

Stress, adaptation and evolution are major concerns in conservation biology. Stresses from pollution, climatic changes, disease etc. may affect population persistence. Further, stress typically occurs when species are placed in captivity. Threatened species are usually managed to conserve their ability to adapt to environmental changes, whilst species in captivity undergo adaptations that are deleterious upon reintroduction into the wild. In model studies using Drosophila melanogaster, we have found that; (a) inbreeding and loss of genetic variation reduced resistance to the stress of disease, (b) extinction rates under inbreeding are elevated by stress, (c) adaptive evolutionary potential in an increasingly stressful environment is reduced in small population, (d) rates of inbreeding are elevated under stressful conditions, (e) genetic adaptation to captivity reduces fitness when populations are reintroduced into the 'wild', and (f) the deleterious effects of adaptation on reintroduction success can be reduced by population fragmentation.     

Regulation of E- and P-cadherin expression correlated with melanocyte migration and diversification.

Melanocytes (Mc) and their progenitors melanoblasts (Mb) are derived from the neural crest and migrate along the dorsolateral pathway to colonize the dermis, the epidermis, and finally the hair matrix. To examine the involvement of cadherins in the migration of Mc lineage cells, we combined flow cytometric analysis of dissociated live cells with immunohistochemical staining of tissue sections to quantify the level of cadherin expression on the surface of Mb/Mc. At 11.5 days postcoitum, Mb are in the dermis and are E-cadherin(-)P-cadherin(-) (E-cad(-)P-cad(-)). During the next 48 h, a 200-fold increase of E-cadherin expression is induced on the surface of Mb prior to their entry into the epidermis, thereby forming a homogeneous E-cad(high)P-cad(-/low) population. The cadherin expression pattern then diversifies, giving rise to three populations, an E-cad(-)P-cad(-) dermal population, E-cad(high)P-cad(low) epidermal population, and E-cad(-)P-cad(med-high) follicular population. In all three populations, the patterns of expression are region-specific, being identical with those of surrounding cells such as keratinocytes and fibroblasts, and are preserved before and after pigmentation. While most of the epidermal Mb/Mc disappear after the neonatal stage in normal mice, forced expression of steel factor in the epidermis of transgenic mice promotes survival of epidermal Mb/Mc, maintaining epidermal-type cadherin expression pattern (E-cad(high)P-cad(low)) throughout the postnatal life. These findings indicate the involvement of extrinsic cues in coordinating the cadherin expression pattern of Mb/Mc and suggest a role for E- and P-cadherins in guiding Mc progenitors to their final destinations.     

Phytochemistry and the systematics and ecology of Loasaceae and Gronoviaceae (Loasales)

A screening for iridoid compounds of 78 of 315 species from all major groups in Gronoviaceae and Loasaceae has been carried out. The results were compared to the systematic concepts in the family and distribution and ecology of the taxa. Iridoids are present in at least some species of all genera screened. Some simple, monomeric compounds (e.g., loganin, sweroside) are found in all major groups of the two families and represent the basic iridoid inventory. Other compounds are restricted to certain taxonomic groups: nine-carbon iridoids (e.g., deutzioside) are restricted to Mentzelia (Loasaceae subfam. Mentzelioideae), hetero-oligomeric iridoids (e.g., tricoloroside methyl ester, acerifolioside) are restricted to two small groups in Loasa (Loasa ser. Macrospermae and ser. Floribundae, Loasaceae subfam. Loasoideae), and oleosides (e.g., 10-hydroxyoleoside dimethyl ester) are restricted to the large genus Caiophora sensu Weigend). The distribution of certain iridoid compounds thus confirms some of the generic limits. Iridoid phytochemistry does not correlate with systematic entities above the generic level nor does it in any way correlate with the morphological evolution of taxa. Conversely, the amount and complexity of iridoid compounds present in taxa correlate positively with the aridity of their habitat and the extent of mammalian herbivore pressure.     

Leucophores and iridophores of Fundulus heteroclitus: Biophysical and ultrastructural properties

Classical light microscopic studies on pigmentation of Fundulus heteroclitus (killifish) indicated that there are three groups of light reflecting cells; one group on the surface of scales reflects white light, while two other deeper groups (the melaniridophores and the stratum argenteum) are iridescent. The results presented here show that: (1) The scale leucophores reflect white light by a Tyndall light-scattering mechanism, by virtue of the presence of randomly oriented organelles of “novel” morphology. (2) The iridophores of the melaniridophores contain stacks of irregularly-spaced, large reflecting platelets which function as an imperfect multiple thin layer interference system. (3) The stratum argenteum consists of a continuous layer(s) of iridophores with reflecting platelets which are so regularly packed as to approach an ideal multiple thin layer interference system. (4) In all three types of light reflecting cells, the dimensions and packing (orientation) of the reflecting organelles satisfactorily account for the chromogenic properties of the cells, including colors as viewed under transmitted, reflected, or polarized light. (5) The spacial relationships between these light reflecting cells and adjoining melanophores are different for each type of light reflecting cell. Furthermore, we propose to replace the term reflecting platelet with refractosome.     

The electronic properties of poly(p-phenylenevinylene) derivatives and their monomers and oligomers

The electronic structures of monomers, oligomers and polymers of poly(p-phenylenevinylene) (PPV) derivatives are calculated and analysed based on density functional theory (DFT) methods. The influences of different substituent groups on the band gaps are discussed. Strong relationships are found between band-gap and bond length alternation (BLA) of polymers, and between band-gap and Wiberg bond index (WBI). Analysis of nuclear independent chemical shift (NICS) reveals that oligomers with similar energy gaps have close values of NICS.     

Treponema denticola cystalysin catalyzes beta-desulfination of L-cysteine sulfinic acid and beta-decarboxylation of L-aspartate and oxalacetate

Pyridoxal 5'-phosphate-dependent cystalysin from Treponema denticola catalyzes the beta-displacement of the beta-substituent from both L-aspartate and L-cysteine sulfinic acid. The steady-state kinetic parameters for beta-desulfination of L-cysteine sulfinic acid, k(cat) and K(m), are 89+/-7 s(-1) and 49+/-9 mM, respectively, whereas those for beta-decarboxylation of L-aspartate are 0.8+/-0.1 s(-1) and 280+/-70 mM. Moreover, cystalysin in the pyridoxamine 5'-phosphate form has also been found to catalyze beta-decarboxylation of oxalacetate as shown by consumption of oxalacetate and a concomitant production of pyruvate. The k(cat) and K(m) of this reaction are 0.15+/-0.01 s(-1) and 13+/-2 mM, respectively. Possible mechanistic and physiological implications are discussed.     

Vernation patterns in Celtidaceae and Ulmaceae (Urticales), and their evolutionary and systematic implications

In two associated families, Celtidaceae and Ulmaceae, vernation pattern (represented by spatial relationships between leaf lamina and stipules, the presence or absence of stipular fusion, lamina orientation, and lamina folding pattern) is consistent within a genus but shows a significant diversity within a family. Six vernation types are distinguishable and tentatively named: 1) Celtis type (Aphananthe, Celtis, Lozanella, Parasponia, Pteroceltis, Trema), 2) Chaetachme type (Chaetachme), 3) Gironniera type (Gironniera), 4) Holoptelea type (Ampelocera, Holoptelea, Phyllostylon), 5) Zelkova type (Hemiptelea, Planera, Zelkova), and 6) Ulmus type (Ulmus). The former three types (found in most of celtidaceous genera) possess free or fused stipules inside of the lamina; in contrast, the latter three types (found in all six ulmaceous genera andAmpelocera) are characterized by having the free stipules outside of the lamina. Within Celtidaceae, Celtis type is probably primitive in having free stipules and an ordinarily oriented lamina; Chaetachme type (with fused, convolute stipules and obliquely oriented laminas) and Gironniera type (with laterally oriented laminas) are the derived. Likewise, within Ulmaceae, both Zelkova and Ulmus types (with laterally oriented laminas) are the derived, while Holoptelea type (with ordinarily oriented laminas) is primitive. Comparisons in vernation pattern suggest the distinctness of Celtidaceae from Ulmaceae and the isolated position ofAmpelocera.