The modern theory of biological evolution: an expanded synthesis

In 1858, two naturalists, Charles Darwin and Alfred Russel Wallace, independently proposed natural selection as the basic mechanism responsible for the origin of new phenotypic variants and, ultimately, new species. A large body of evidence for this hypothesis was published in Darwins Origin of Species one year later, the appearance of which provoked other leading scientists like August Weismann to adopt and amplify Darwins perspective. Weismanns neo-Darwinian theory of evolution was further elaborated, most notably in a series of books by Theodosius Dobzhansky, Ernst Mayr, Julian Huxley and others. In this article we first summarize the history of life on Earth and provide recent evidence demonstrating that Darwins dilemma (the apparent missing Precambrian record of life) has been resolved. Next, the historical development and structure of the modern synthesis is described within the context of the following topics: paleobiology and rates of evolution, mass extinctions and species selection, macroevolution and punctuated equilibrium, sexual reproduction and recombination, sexual selection and altruism, endosymbiosis and eukaryotic cell evolution, evolutionary developmental biology, phenotypic plasticity, epigenetic inheritance and molecular evolution, experimental bacterial evolution, and computer simulations (in silico evolution of digital organisms). In addition, we discuss the expansion of the modern synthesis, embracing all branches of scientific disciplines. It is concluded that the basic tenets of the synthetic theory have survived, but in modified form. These sub-theories require continued elaboration, particularly in light of molecular biology, to answer open-ended questions concerning the mechanisms of evolution in all five kingdoms of life.Dedicated to Prof. Dr. Dr. hc mult. Ernst Mayr on the occasion of his 100th birthdayThis revised version was published online in March 2004, with corrections to the caption of Figure 6.     

Hopanoid Biosynthesis and Function in Bacteria

   Hopanoids are pentacyclic triterpenoid lipids occurring in bacteria. They are synthesized from isopentenyl units which are formed in a new biosynthetic route leading to isopentenyl diphosphate. Six C₅ units are joined to form squalene, the immediate precursor in hopanoid synthesis. In a highly complex cyclization reaction that shares considerable similarities with that of oxidosqualene to sterols, the hopane skeleton is formed from squalene by the squalene-hopene cyclase. Recent elucidation of the X-ray structure of this membrane-bound cyclase has shed some light on the properties of this unusual enzyme. The active site is located in a cavity within the enzyme. The squalene substrate diffuses through a channel structure from the membrane into this cavity and is there transformed into hopene. Polar side chains are attached to hopene resulting in the amphiphilic molecular structure of many hopanoids. These hopanoids are membrane components involved in regulating membrane fluidity and stability. However, the many structural variants of hopanoids indicate that they may have other interesting but as yet unknown functions.     

Macroinvertebrate communities in agriculturally impacted southern Illinois streams: patterns with riparian vegetation, water quality, and in-stream habitat quality

Relationships between riparian land cover, in-stream habitat, water chemistry, and macroinvertebrates were examined in headwater streams draining an agricultural region of Illinois. Macroinvertebrates and organic matter were collected monthly for one year from three intensively monitored streams with a gradient of riparian forest cover (6, 22, and 31% of riparian area). Bioassessments and physical habitat analyses were also performed in these three streams and 12 other nearby headwater streams. The intensively monitored site with the least riparian forest cover had significantly greater percent silt substrates than the sites with medium and high forest cover, and significantly higher very fine organics in substrates than the medium and high forested sites. Macroinvertebrates were abundant in all streams, but communities reflected degraded conditions; noninsect groups, mostly oligochaetes and copepods, dominated density and oligochaetes and mollusks, mostly Sphaerium and Physella, dominated biomass. Of insects, dipterans, mostly Chironomidae, dominated density and dipterans and coleopterans were important contributors to biomass. Collector-gatherers dominated functional structure in all three intensively monitored sites, indicating that functional structure metrics may not be appropriate for assessing these systems. The intensively monitored site with lowest riparian forest cover had significantly greater macroinvertebrate density and biomass, but lowest insect density and biomass. Density and biomass of active collector-filterers (mostly Sphaerium) decreased with increasing riparian forest. Hilsenhoff scores from all 15 sites were significantly correlated with in-stream habitat scores, percent riparian forest, and orthophosphate concentrations, and multiple regression indicated that in-stream habitat was the primary factor influencing biotic integrity. Our results show that these "drainage ditches" harbor abundant macroinvertebrates that are typical of degraded conditions, but that they can reflect gradients of conditions in and around these streams.     

Assessing Landscape Condition Relative to Water Resources in the Western United States: A Strategic Approach

The Environmental Monitoring and Assessment Program (EMAP) is proposing an ambitious agenda to assess the status of streams and estuaries in a 12-State area of the western United States by the end of 2003. Additionally, EMAP is proposing to access landscape conditions as they relate to stream and estuary conditions across the west. The goal of this landscape project is to develop a landscape model that can be used to identify the relative risks of streams and estuaries to potential declines due to watershed-scale, landscape conditions across the west. To do so, requires an understanding of quantitative relationships between landscape composition and pattern metrics and parameters of stream and estuary conditions. This paper describes a strategic approach for evaluating the degree to which landscape composition and pattern influence stream and estuary condition, and the development and implementation of a spatially-distributed, landscape analysis approach.