Review Article
Chromosome evolution in the Salmonidae (Pisces): an update
- RUTH PHILLIPS, PETR RÁB
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- Published online by Cambridge University Press:
- 27 March 2001, pp. 1-25
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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.
Sources and bioavailability of phosphorus fractions in freshwaters: a British perspective
- C. S. REYNOLDS, P. S. DAVIES
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- 27 March 2001, pp. 27-64
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This paper seeks a perspective on the forms of phosphorus which promote aquatic eutrophication, with the particular quest of establishing their sources. A short background traces the development of understanding of nutrient enrichment and the suppositions about the relative contributions of agriculture, sewage and detergent residues. Most aquatic systems, and their primary producers, are naturally deficient in biologically-available phosphorus. Aquatic plants have evolved very efficient phosphorus uptake mechanisms. The biomass responses to an increase in the supply of phosphorus are stoichiometrically predictable. The most bioavailable forms of phosphorus are in solution, as orthophosphate ions, or are readily soluble or elutable from loose combinations. Ready bioavailability coincides well with what is measurable as molybdate-reactive (MRP) or soluble-reactive phosphorus (SRP). Most other forms, including phosphates of the alkaline earth metals, aluminium and iron are scarcely available at all. Orthophosphate ions sorbed to metal oxides and hydroxides are normally not biologically available either, except through weak dissociation (‘desorption’). The production of alkaline phosphatase provides organisms with an additional mechanism for accelerating the sequestration of phosphate from organic compounds. Bioavailable phosphate is liberated when redox- or alkali-sensitive metal hydroxides dissolve but these processes are minor contributors to the biological responses to nutrient enrichment.
Most of the familiar eutrophication is attributable to the widespread application of secondary sewage treatment methods to the wastes emanating from a burgeoning and increasingly urbanised human population. The use of polyphosphate-based detergents, now in decline, has contributed to the problem. In aquatic systems, the additional phosphorus raises the biological supportive capacity, sometimes to the capacity of the next limiting factor (carbon, light, hydraulic retention or of another nutrient). At high orthophosphate loadings, the straight stoichiometric yield relationship between biomass yield and phosphorus availability is lost.
Movements of phosphorus and its recycling within aquatic systems do not prevent the slow gravitation of phosphorus to the bottom substrata. The phosphorus retentivity of sediments depends upon their chemical composition. While oxide-hydroxide binding capacity in the surface sediments persists, they act as a sink for phosphorus and a control on further cycling. Iron-rich and clay-rich sediments perform best in these conditions; calcareous sediments least so. Eutrophication may lead to the exhaustion of sediment P-binding capacity. Non-sorbed phosphate is readily recyclable if primary producers have access to it. Recycling is most rapid in shallow waters (where sediment disturbance, by flow, by wind action and through bioturbation, is frequent) and least in deep ventilated sediments.
The contributions of phosphorus from catchments are assessed. The slow rate of weathering of (mostly apatitic) minerals, the role of chemical binding in soils and the incorporation and retentivity by forested terrestrial ecosystems each contribute to the minimisation of phosphorus leakage to drainage waters. Palaeolimnological and experimental evidence confirms that clearance of land and ploughing its surface weakens the phosphorus retentivity of catchments. The phosphorus transferred from arable land to drainage remains dominated by sorbed fractions which are scarcely bioavailable. Some forms of intensive market gardening or concentrated stock rearing may mobilise phosphates to drainage but it is deduced that drainage from agricultural land is not commonly a major source of readily bioavailable phosphorus in water. Careful budgeting of the phosphates in run-off from over-fertilised soils may nevertheless show that a proportionately small loss of bioavailable phosphorus can still be highly significant in promoting aquatic plant production. The bioavailable-phosphorus (BAP) load achieving the OECD threshold of lake eutrophy (35 mg P m−3) is calculated to be equivalent to a terrestrial loss rate of approximately 17.5 kg BAP km−2 year−1), or only 1–2% of a typical fertiliser application. The output is shown to be comparable with the P yield from secondary treatment of the sewage produced by a resident population of 30–44 persons km−2. With tertiary treatment, the equivalence is with approximately 200 persons km−2.
Coincidence, coevolution, or causation? DNA content, cell size, and the C-value enigma
- T. RYAN GREGORY
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- 27 March 2001, pp. 65-101
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Variation in DNA content has been largely ignored as a factor in evolution, particularly following the advent of sequence-based approaches to genomic analysis. The significant genome size diversity among organisms (more than 200 000-fold among eukaryotes) bears no relationship to organismal complexity and both the origins and reasons for the clearly non-random distribution of this variation remain unclear. Several theories have been proposed to explain this ‘C-value enigma’ (heretofore known as the ‘C-value paradox’), each of which can be described as either a ‘mutation pressure’ or ‘optimal DNA’ theory. Mutation pressure theories consider the large portion of non-coding DNA in eukaryotic genomes as either ‘junk’ or ‘selfish’ DNA and are important primarily in considerations of the origin of secondary DNA. Optimal DNA theories differ from mutation pressure theories by emphasizing the strong link between DNA content and cell and nuclear volumes. While mutation pressure theories generally explain this association with cell size as coincidental, the nucleoskeletal theory proposes a coevolutionary interaction between nuclear and cell volume, with DNA content adjusted adaptively following shifts in cell size. Each of these approaches to the C-value enigma is problematic for a variety of reasons and the preponderance of the available evidence instead favours the nucleotypic theory which postulates a causal link between bulk DNA amount and cell volume. Under this view, variation in DNA content is under direct selection via its impacts on cellular and organismal parameters. Until now, no satisfactory mechanism has been presented to explain this nucleotypic effect. However, recent advances in the study of cell cycle regulation suggest a possible ‘gene–nucleus interaction model’ which may account for it. The present article provides a detailed review of the debate surrounding the C-value enigma, the various theories proposed to explain it, and the evidence in favour of a causal connection between DNA content and cell size. In addition, a new model of nucleotypic influence is developed, along with suggestions for further empirical investigation. Finally, some evolutionary implications of genome size diversity are considered, and a broadening of the traditional ‘biological hierarchy’ is recommended.
Ethylene signal perception and transduction: multiple paradigms?
- M. A. HALL, I. E. MOSHKOV, G. V. NOVIKOVA, L. A. J. MUR, A. R. SMITH
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- 27 March 2001, pp. 103-128
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Current progress on the mechanisms of ethylene signal perception and transduction are reviewed with an emphasis on reconciling data from molecular genetics and from biochemical approaches. It is proposed that there exist two or more interacting transduction pathways.
From arctic lemmings to adaptive dynamics: Charles Elton's legacy in population ecology
- JAN LINDSTRÖM, ESA RANTA, HANNA KOKKO, PER LUNDBERG, VEIJO KAITALA
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- 27 March 2001, pp. 129-158
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We shall examine the impact of Charles S. Elton's 1924 article on periodic fluctuations in animal populations on the development of modern population ecology. We argue that his impact has been substantial and that during the past 75 years of research on multi-annual periodic fluctuations in numbers of voles, lemmings, hares, lynx and game animals he has contributed much to the contemporary understanding of the causes and consequences of population regulation. Elton was convinced that the cause of the regular fluctuations was climatic variation. To support this conclusion, he examined long-term population data then available. Despite his firm belief in a climatic cause of the self-repeating periodic dynamics which many species display, Elton was insightful and far-sighted enough to outline many of the other hypotheses since put forward as an explanation for the enigmatic long-term dynamics of some animal populations. An interesting, but largely neglected aspect in Elton's paper is that it ends with speculation regarding the evolutionary consequences of periodic population fluctuations. The modern understanding of these issues will also be scrutinised here. In population ecology, Elton's 1924 paper has spawned a whole industry of research on populations displaying multi-annual periodicity. Despite the efforts of numerous research teams and individuals focusing on the origins of multi-annual population cycles, and despite the early availability of different explanatory hypotheses, we are still lacking rigorous tests of some of these hypotheses and, consequently, a consensus of the causes of periodic fluctuations in animal populations. Although Elton would have been happy to see so much effort spent on cyclic populations, we also argue that it is unfortunate if this focus on a special case of population dynamics should distract our attention from more general problems in population and community dynamics.
Correction
Author's errata
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- Published online by Cambridge University Press:
- 27 March 2001, p. 159
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The following corrections are made to the paper ‘Trophic-dynamic considerations in relating species diversity to ecosystem resilience’ by Kris H. Johnson, which was published in Biological Reviews75(3) (2000), 347–376.
Erratum 1:
p. 361: the first complete sentence following equation (20) should read as follows: ‘Thus, Φ is expected to be inversely proportionate to the amount of community-wide trophic specialization characterizing the system and to C from equation (8), and directly proportionate to both R and Γ.’
Erratum 2:
p. 363: the third full sentence from the right-hand column (line 10) should begin as follows: ‘Φ which is directly proportionate to R....’