Origin and relationships of the myco-heterotrophic liverwort Cryptothallus mirabilis Malmb. (Metzgeriales, Marchantiophyta)

Heterotrophic life histories have evolved independently numerous times in the angiosperms. In non-vascular embryophytes, heterotrophy is known only in the genus Cryptothallus . Cryptothallus mirabilis obtains photosynthates indirectly from a host tree via a basidiomycete that is simultaneously ectomycorrhizal on the host, a strategy known as myco-heterotrophism. This simple thalloid liverwort was initially described as an albino variant of Aneura pinguis , and the literature varies on whether it should be considered as such, as a distinct species of Aneura , or as a separate genus. Here, the relationships of C. mirabilis within the family Aneuraceae are reconstructed using DNA sequence data from the chloroplast ( rps 4, rps 14, atp B- rbc L spacer, trn G), mitochondrial ( trn S), and nuclear (26S and ITS) genomes. Several allopatric populations of C. mirabilis and of both sympatric species of Aneura (i.e. A. pinguis and A. maxima ) were included. Cryptothallus mirabilis is resolved as having a single origin from within Aneura , and hence the myco-heterotrophic liverwort should be considered as a distinct species of Aneura , rather than an autonomous genus. The fungal symbiont of the photosynthetic A. pinguis is of the same genus, Tulasnella , as that of C. mirabilis , suggesting that the heterotrophic life strategy might have evolved from a pre-existing symbiosis.  © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society , 2008, 156 , 1–12.     

The evolution of very-low-calorie diets: an update and meta-analysis

Objective: Very‐low‐calorie diets (VLCDs), providing <800 kcal/d, have been used since the 1970s to induce rapid weight loss. Previous reviews of the literature have disagreed concerning the relative efficacy of VLCDs vs. conventional low‐calorie diets (LCDs) for achieving long‐term weight loss. Research Methods and Procedures: We sought to update findings on the clinical use, safety, and efficacy of VLCDs and to perform a meta‐analysis of randomized trials that compared the long‐term efficacy of LCDs and VLCDs. Original research articles were retrieved by a Medline search and from prior reviews of VLCDs. Trials were included only if they were randomized comparisons of LCDs and VLCDs and included a follow‐up assessment at least 1 year after maximum weight loss. Data were abstracted by both authors regarding: duration of VLCD, total length of treatment, attrition, short‐ and long‐term weight loss, changes in weight‐related comorbidities, and adverse effects. Results: Six randomized trials were found that met inclusion criteria. VLCDs, compared with LCDs, induced significantly greater short‐term weight losses (16.1 ± 1.6% vs. 9.7 ± 2.4% of initial weight, respectively; p = 0.0001) but similar long‐term losses (6.3 ± 3.2% vs. 5.0 ± 4.0%, respectively; p > 0.2). Attrition was similar with VLCD and LCD regimens. Discussion: VLCDs did not produce greater long‐term weight losses than LCDs. In the United States, the use of liquid meal replacements as part of a 1000 to 1500 kcal/d diet may provide an effective and less expensive alternative to VLCDs. In Europe, VLCDs are used with less intensive medical supervision than in the United States, which reduces the cost of this approach.     

Mycoheterotrophic interactions are not limited to a narrow phylogenetic range of arbuscular mycorrhizal fungi

The majority of achlorophyllous mycoheterotrophic plant species associate with arbuscular mycorrhizal fungi (AMF). Previous studies have shown that some species are highly specialized towards narrow lineages of AMF and have suggested that only particular lineages of these fungi are targeted by mycoheterotrophic plants. To test this hypothesis, we analyzed all available partial SSU sequences of AMF associated with mycoheterotrophic plants including data from 13 additional specimens from French Guiana, Gabon and Australia. Sequences were assigned to 'virtual taxa' (VT) according to the MaarjAM database. We found that 20% of all known Glomeromycota VT are involved in mycoheterotrophic interactions and the majority of associations involve Glomeraceae (Glomus Group A) fungi. While some mycoheterotrophic plant species have been found growing with only a single VT, many species are able to associate with a wide range of AMF. We calculated significant phylogenetic clustering of Glomeromycota VT involved in mycoheterotrophic interactions, suggesting that associations between mycoheterotrophic plants and AMF are influenced by the phylogenetic relationships of the fungi. Our results demonstrate that many lineages of AMF are prone to exploitation by mycoheterotrophic plants. However, mycoheterotrophs from different plant lineages and different geographical regions tend to be dependent on lineages of AMF that are phylogenetically related.     

Directed evolution of industrial enzymes: an update

The use of enzymes in industrial processes can often eliminate the use of high temperatures, organic solvents and extremes of pH, while at the same time offering increased reaction specificity, product purity and reduced environmental impact. The growing use of industrial enzymes is dependent on constant innovation to improve performance and reduce cost. This innovation is driven by a rapidly increasing database of natural enzyme diversity, recombinant DNA and fermentation technologies that allow this diversity to be produced at low cost, and protein modification tools that enable enzymes to be tuned to fit into the industrial marketplace.     

Pteridophyte fungal associations: Current knowledge and future perspectives

Current understanding of the nature and function of fungal associations in pteridophytes is surprisingly patchy given their key evolutionary position, current research foci on other early-branching plant clades, and major efforts at unravelling mycorrhizal evolution and the mechanisms underlying this key interaction between plants and fungi. Here we provide a critical review of current knowledge of fungal associations across pteridophytes and consider future directions making recommendations along the way. From a comprehensive survey of the literature, a confused picture emerges: suggestions that members of the Lycopsida harbour Basidiomycota fungi contrast sharply with extensive cytological and recent molecular evidence pointing to exclusively Glomeromycota and/or Mucoromycotina associations in this group. Similarly, reports of dark septate, assumingly ascomycetous, hyphae in a range of pteridophytes, advocating a mutualistic relationship, are not backed by functional evidence and the fact that the fungus invariably occupies dead host tissue points to saprotrophy and not mutualism. The best conclusion that can be reached based on current evidence is that the fungal symbionts of pteridophytes belong to the two fungal lineages Mucoromycotina and Glomeromycota. Do symbiotic fungi and host pteridophytes engage in mutually beneficial partnerships? To date, only two pioneering studies have addressed this key question demonstrating reciprocal exchange of nutrients between the sporophytes of Ophioglossum vulgatum and Osmunda regalis and their fungal symbionts. There is a pressing need for more functional investigations also extending to the gametophyte generation and coupled with in vitro isolation and resynthesis studies to unravel the effect of the fungi on their host.     

Glomeromycotean associations in liverworts: a molecular, cellular, and taxonomic analysis

Liverworts form endophytic associations with fungi that mirror mycorrhizal associations in tracheophytes. Here we report a worldwide survey of liverwort associations with glomeromycotean fungi (GAs), together with a comparative molecular and cellular analysis in representative species. Liverwort GAs are circumscribed by a basal assemblage embracing the Haplomitriopsida, the Marchantiopsida (except a few mostly derived clades), and part of the Metzgeriidae. Fungal endophytes from Haplomitrium, Conocephalum, Fossombronia, and Pellia were related to Glomus Group A, while the endophyte from Monoclea was related to Acaulospora. An isolate of G. mosseae colonized axenic thalli of Conocephalum, producing an association similar to that in the wild. Fungal colonization in marchantialean liverworts suppressed cell wall autofluorescence and elicited the deposition of a new wall layer that specifically bound the monoclonal antibody CCRC-M1 against fucosylated side groups associated with xyloglucan and rhamnogalacturonan I. The interfacial material covering the intracellular fungus contained the same epitopes present in host cell walls. The taxonomic distribution and cytology of liverwort GAs suggest an ancient origin and multiple more recent losses, but the occurence in widely separated liverwort taxa of fungi related to glomeromycotean lineages that form arbuscular mycorrhizas in tracheophytes, notably the Glomus Group A, is better explained by host shifting from tracheophytes to liverworts.     

Chromosome evolution in the Salmonidae (Pisces): an update

The karyotypes of salmonid fishes including taxa in the three subfamilies Coregoninae, Thymallinae and Salmoninae are described. This review is an update of the (Hartley, 1987) review of the chromosomes of salmonid fishes. As described in the previous review, the karyotypes of salmonid fishes fall into two main categories based on chromosome numbers: the type A karyotypes have diploid numbers close to 80 with approximately 100 chromosome arms (2n = 80, NF = 100), and the type B karyotypes have diploid numbers close to 60 with approximately 100 chromosome arms (2n = 60, NF = 100). In this paper we have proposed additional sub categories based on variation in the number of chromosome arms: the A' type with NF = 110-120, the A" type with NF greater than 140, and the B' type with NF less than 80. Two modes of chromosome evolution are found in the salmonids: in the Coregoninae and the Salmoninae the chromosomes have evolved by centric fusions of the Robertsonian type decreasing chromosome numbers (2n) while retaining chromosome arm numbers (NF) close to that found in the hypothetical tetraploid ancestor so that most extant taxa have either type A or type B karyotypes. In the Thymallinae, the chromosomes have evolved by inversions so that chromosome arm numbers (NF) have increased but chromosome numbers (2n) close to the karyotype of the hypothetical tetraploid ancestor have been retained and all taxa have type A' karyotypes. Most of the taxa with type B karyotypes in the Coregoninae and Salmoninae are members of the genus Oncorhynchus, although at least one example of type B karyotypes is found in all of the other genera. These taxa either have an anadromous life history or are found in specialized lacustrine environments. Selection for increases or decreases in genetic recombination as proposed by Qumsiyeh, 1994 could have been involved in the evolution of chromosome number in salmonid fishes.     

Brassicales: an update on chromosomal evolution and ancient polyploidy

Brassicales comprise 17 families, c. 400 genera and more than 4600 species. Despite the mustard family (crucifers, Brassicaceae) continuing to be the subject of intensive research, the remaining 16 families are largely under studied. Here I summarize the available data on chromosome number and genome size variation across Brassicales in the context of a robust phylogenetic framework. This analysis has revealed extensive knowledge gaps in karyological data for non-crucifer and species-rich families in particular (i.e., Capparaceae, Cleomaceae, Resedaceae and Tropaeolaceae). A parsimonious interpretation of the combined chromosomal and phylogenetic data set suggests that the ancestral pre-Brassicales genome had 9 or 14 chromosome pairs, later multiplied by the At-β (beta) whole-genome duplication (WGD) to n?=?18 or 28. This WGD was followed by post-polyploid diploidization marked by diversification to 12 or 13 families and independent decreases in chromosome numbers. Family-specific WGDs are proposed to precede the diversification of Capparaceae, Resedaceae and Tropaeolaceae.     

The evolution of insect/vertebrate associations

The evolution of close vertebrate associations has occurred in seven orders of insects, resulting in a great diversity of interactions which range from commensalism to true parasitism. The evolution of each taxon of vertebrate associates is discussed in turn, some new ideas on the development of certain groups are presented and, on a broader scale, a general model for the evolution of ectoparasitic insects is proposed. It argues that all vertebrate associates have evolved along one of two macroevolutionary pathways which differ only in the sequencing of adaptations facilitating host association and host feeding. These pathways lead to parasite types which differ greatly in their life history and intimacy of host association.
Some microevolutionary processes influencing the diversification of ectoparasites are discussed, in particular the process of insect/vertebrate coevolution and the forms this may take. Host specificity, one consequence of coevolution, is recognised as an important factor influencing the structure of ectoparasite communities, and a hypothesis is presented that competition between ectoparasite species, mediated by host defensive responses, is also important in determining community structure.     

Pattern and process in the evolution of insect-plant associations: Yponomeuta as an example

Phylogenetic studies are increasing our understanding of the evolution of associations between phytophagous insects and their host plants. Sequential evolution, i.e. the shift of insect herbivores onto pre-existing plant species, appears to be much more common than coevolution, where reciprocal selection between interacting insects and plants is thought to induce chemical diversification and resistance in plants and food specialization in insects.Extreme host specificity is common in phytophagous insects and future studies are likely to reveal even more specialization. Hypotheses that assume that food specialists have selective advantages over generalists do not seem to provide a general explanation for the ubiquity of specialist insect herbivores. Specialists are probably committed to remain so, because they have little evolutionary opportunity to reverse the process due to genetically determined constraints on the evolution of their physiology or nervous system. The same constraints might result in phylogenetic conservatism, i.e. the frequent association of related insect herbivores with related plants. Current phylogenetic evidence, however, indicates that there is no intrinsic direction to the evolution of specialization.Historical aspects of insect-host plant associations will be illustrated with the small ermine moth genus Yponomeuta. Small ermine moths show an ancestral host association with the family Celastraceae. The genus seems to be committed to specialization per se rather than to a particular group of plants. Whatever host shift they have made in their evolutionary past (onto Rosaceae, Crassulaceae, and Salicaceae), they remain monophagous. The oligophagous Y. padellus is the only exception. This species might comprise a mosaic of genetically divergent host-associated populations.     

Co-culture of arbuscular mycorrhiza-like fungi (Piriformospora indica and Sebacina vermifera) with plant cells of Linum album for enhanced production of podophyllotoxins: a first report

Cell suspension cultures of Linum album were developed from internode portions of in vitro germinated plant in Gamborg's B5 medium supplemented with 0.4 mg naphthalene acetic acid/l. The highest biomass was 8.5 g/l with podophyllotoxin and 6-methoxypodophyllotoxin at 29 and 1.9 mg/l, respectively after 12 d cultivation. Co-cultures of L. album cells with axenically cultivable arbuscular mycorrhiza-like fungi, Piriformospora indica and Sebacina vermifera, were established for the first time. These enhanced podophyllotoxin and 6-methoxypodophyllotoxin production by about four- and eight-fold, respectively, along with a 20% increase in biomass compared to the control cultures.     

The developmental evolution of avian digit homology: An update

The identity of avian digits has been unresolved since the beginning of evolutionary morphology in the mid-19th century, i.e. as soon as questions of phylogenetic homology have been raised. The main source of concern is the persistent discrepancy between anatomical/paleontological and embryological evidence over the identity of avian digits. In this paper, recent evidence pertaining to the question of avian digit homology is reviewed and the various ideas of how to resolve the disagreement among developmental and phylogenetic evidence are evaluated. Paleontological evidence unequivocally supports the hypothesis that the fully formed digits of maniraptoran theropods are digits DI, DII, and DIII, because the phylogenetic position of Herrerasaurus is resolved, even when hand characters are excluded from the analysis. Regarding the developmental origin of the three digits of the avian hand the discovery of an anterior digit condensation in the limb bud of chickens and ostriches conclusively shows that these three digits are developing from condensations CII, CIII, and CIV. The existence of this additional anterior condensation has been confirmed in four different labs, using four different methods: Alcian blue staining, PNA affinity histochemistry, micro-capillary regression and Sox9 expression. Finally, recent evidence shows that the digit developing from condensation CII has a Hox gene expression pattern that is found in digit DI of mice forelimb and chick hind limbs. The sum of these data supports the idea that digit identity has shifted relative to the location of condensations, known as Frame Shift Hypothesis, such that condensation CII develops into digit DI and condensation CIII develops into digit DII, etc. A review of the literature on the digit identity of the Italian Three-toed Skink or Luscengola (Chalcides chalcides), shows that digit identity frame shifts may not be limited to the bird hand but may be characteristic of “adaptive” digit reduction in amniotes (sensu Steiner, H., Anders, G., 1946. Zur Frage der Entstehung von Rudimenten. Die Reduktion der Gliedmassen von Chalcides tridactylus Laur. Rev. Suisse Zool. 53, 537–546) in general. In this mode of evolution two digits are lost, in the course of the adaptation of the three anterior digits to a function that does not require the two posterior digits. This evidence suggests that the evolution of digits in tetrapods can proceed at least on two distinct levels of integration, the level of digit condensations and that of adult digits.     

The new mycobacteria: an update

The continuous evolution of mycobacterial taxonomy may represent a source of confusion for laboratories and clinicians. Apart from the obvious pathogenic strains of the Mycobacterium tuberculosis complex, Mycobacterium leprae and Mycobacterium ulcerans, the role of other mycobacteria may be associated with varying conditions ranging from contamination to specific disease processes. Of the more than 120 mycobacterial species recognized currently, very few have not been reported as pathogenic in humans or animals. Although the attempt to keep pace with the steadily increasing number of mycobacterial species seems hopeless, a careful review of the recent literature relevant to the newly described species may be advantageous. The aim of this present update is to provide epidemiological and clinical information along with major phenotypic and genotypic characteristics of the species described in the last 3 years.     

The freemartin syndrome: an update

The freemartin condition represents the most frequent form of intersexuality found in cattle, and occasionally other species. This review considers the current state of knowledge of freemartin biology, incidence, experimental models, diagnosis, uses for freemartins in cattle herds, occurrence in non-bovine species, effects on the male, and highlights potential new research areas. Freemartins arise when vascular connections form between the placentae of developing heterosexual twin foeti, XX/XY chimerism develops, and ultimately there is masculinisation of the female tubular reproductive tract to varying degrees. With twinning rates in Holstein cows increasing, there will be greater economic importance to establish early diagnosis of the freemartin and the detection of the less common single born freemartin. New diagnostic methods based on the detection of Y-chromosome DNA segments by polymerase chain reaction (PCR) show improved assay sensitivity and efficiency over karyotyping and clinical examination. The implications for the chimeric male animal born co-twin to the freemartin are contentious as to whether fertility is affected; if germ cell chimerism does indeed occur; and, if there are any real effects on the sex ratio of offspring produced. In beef cattle, the freemartin carcass has similar characteristics to normal herdmates. Hormonal treatment of freemartins for use as oestrous detectors has been used to obtain salvage value. The biology of freemartin sheep has recently been studied in detail, and the condition may be increasing in prevalence with the introduction of high fecundity genes into flocks. Potential new research areas are discussed, such as detection of foetal DNA in maternal circulation for prenatal diagnosis and investigation of the anti-tumour properties of Mullerian inhibiting substance (MIS). The freemartin syndrome will always be a limiting factor in cattle and to a lesser extent in sheep production systems that have the goal to produce multiple reproductively normal female offspring from a single dam without using sex predetermination.     

Role of microorganisms in the evolution of animals and plants: the hologenome theory of evolution

We present here the hologenome theory of evolution, which considers the holobiont (the animal or plant with all of its associated microorganisms) as a unit of selection in evolution. The hologenome is defined as the sum of the genetic information of the host and its microbiota. The theory is based on four generalizations: (1) All animals and plants establish symbiotic relationships with microorganisms. (2) Symbiotic microorganisms are transmitted between generations. (3) The association between host and symbionts affects the fitness of the holobiont within its environment. (4) Variation in the hologenome can be brought about by changes in either the host or the microbiota genomes; under environmental stress, the symbiotic microbial community can change rapidly. These points taken together suggest that the genetic wealth of diverse microbial symbionts can play an important role both in adaptation and in evolution of higher organisms. During periods of rapid changes in the environment, the diverse microbial symbiont community can aid the holobiont in surviving, multiplying and buying the time necessary for the host genome to evolve. The distinguishing feature of the hologenome theory is that it considers all of the diverse microbiota associated with the animal or the plant as part of the evolving holobiont. Thus, the hologenome theory fits within the framework of the 'superorganism' proposed by Wilson and Sober.     

The evolution of actinorhizal symbioses: Evidence for multiple origins of the symbiotic association

According to morphologically based classification systems, actinorhizal plants, engaged in nitrogen-fixing symbioses with Frankia bacteria, are considered to be only distantly related. However, recent phylogenetic analyses of seed plants based on chloroplast rbcL gene sequences have suggested closer relationships among actinorhizal plants. A more thorough sampling of chloroplast rbcL gene sequences from actinorhizal plants and their nonsymbiotic close relatives was conducted in an effort to better understand the phylogenetic relationships of these plants, and ultimately, to assess the homology of the different actinorhizal symbioses. Sequence data from 70 taxa were analyzed using parsimony analysis. Strict consensus trees based on 24 equally parsimonious trees revealed evolutionary divergence between groups of actinorhizal species suggesting that not all symbioses are homologous. The arrangement of actinorhizal species, interspersed with nonactinorhizal taxa, is suggestive of multiple origins of the actinorhizal symbiosis. Morphological and anatomical characteristics of nodules from different actinorhizal hosts were mapped onto the rbclL-based consensus tree to further assess homology among rbcL-based actinorhizal groups. The morphological and anatomical features provide additional support for the rbcL-based groupings, and thus, together, suggest that actinorhizal symbioses have originated more than once in evolutionary history.