Freshwater Phytoplankton Ecology: the British Contribution in Retrospect

1 Article

Freshwater phytoplankton ecology: the British contribution in retrospect

J.F. Talling Hawthorn View, The Pines, Bongate, Appleby, Cumbria CA16 6HR, UK. Email: [email protected]

Received 13 October 2011; accepted 23 December 2011; published 1 June 2012

Abstract

The ecology of the freshwater phytoplankton – communities of freely-dispersed, plant-functioning microbes – has attracted attention in Britain for over 100 years. Here an outline history is given, with reference to persons and places, types of water-body involved, and the approaches adopted. The last include taxonomic resolution and composition with range of morphotypes; physiology of growth, photosynthesis and nutrition in relation to environmental variables; population sampling, dynamics and controlling factors; experimental mesocosms; and synecological aspects of communities. Reductionist and autecological approaches have been accompanied by more abstract generalisation, including phytoplankton viewed as a model community.

Keywords: algal periodicity; chytrid parasitism; clear-water phase; grazing; models; nutrient budget; phytoplankton; population dynamics; seasonality; water-bloom.

Introduction international scope can be found in the works of Lund (1965), Round (1981), Harris (1986), Fogg & Thake (1987), National insularity is not acceptable in scientific thinking, Reynolds (1984, 2006) and Talling (2002). even when there is a literal geographical basis. But for the Readers – especially from outside Britain – might note purpose of historical survey, a limitation of scope has some that the work surveyed has been contributed by general advantages – for coverage of detail, of significant links of naturalists supported by their local societies, university disciplines through localised individual connections, and academics, and the staff-members of state-assisted of expansion in approaches from locally based associations or -related institutions like the Freshwater Biological of individuals, facilities and environment. Here such Association, Nature Conservancy and later regional survey is applied to the development of basic work on derivatives, the Centre for Ecology and Hydrology the ecology of freshwater phytoplankton. References are of the Natural Environment Research Council, and examples only. They are restricted to work carried out in the Department of Agriculture Northern Ireland with the British Isles and some derived projects overseas, and derivatives. Field stations have played a large part. to mainly factual rather than conceptual issues. Historic Issues with water supply involved the Water Research relations with other freshwater science (limnology) in Association and local water authorities, such as the Britain are considered by Talling (2004). Surveys of Metropolitan Water Board (MWB) and its successors which

DOI: 10.1608/FRJ-5.1.453 Freshwater Reviews (2012) 5, pp. 1-20 © Freshwater Biological Association 2012 2 Talling, J.F. faced problems from phytoplankton growth in nutrient-rich English Lakes (West & West, 1912) and Fritsch with his Thames river water stored in reservoirs near London. co-worker Florence Rich the more mixed communities (‘heleoplankton’) of ponds (Fritsch & Rich, 1913). Algal occurrence, communities and Although algal periodicity later attracted others periodicity (Griffiths, 1923; Southern & Gardiner, 1926; Lind, 1938), limitations of sampling and counting before 1940 led to Early interest centred on the visible results of water-blooms somewhat primitive measures of population dynamics (Griffiths, 1939) and of sampling in nature – originally and interpretations of regulating factors. Thereafter mainly by fine plankton nets, which did not retain the improvements followed, many introduced by Fritsch’s smaller forms known later as nanoplankton, ‘µ-algae’ former student J.W.G. Lund (Fig. 1e) (also a skilled algal and picoplankton. The observations led naturally to taxonomist) and his colleagues (Lund & Talling, 1958), taxonomic resolution of species, such as the common including counts using iodine-sedimentation and the diatom Asterionella formosa obtained by A.H. Hassall from inverted microscope (Lund et al., 1958), a novel means of a London reservoir. Notable naturalists included W. West integrated sampling by vertical plastic tube (Lund, 1949), (Fig. 1a), originally a Leeds and Bradford pharmacist, biomass assessment by chlorophyll a, and attention to whose son G.S. West (Fig. 1b) graduated from Cambridge the smallest forms (Lund, 1961; Bailey-Watts et al., 1968; University and contributed richly, with his father or singly, Vincent, 1981; Happey-Wood et al., 1988). Biomass to the recognition of species in Britain (West & West, 1901, assessment by chlorophyll a developed rather late in 1905, 1906a, b; G.S. West, 1904) and elsewhere, including a Britain, partly due to a traditional reluctance to move comparative account of the phytoplankton of three major from microscopy to chemical methods and partly from African lakes (G.S. West, 1907). He also considered seasonal experience of incomplete chlorophyll extraction from algae changes of phytoplankton in an Australian reservoir near of London reservoirs where the method was tested by Melbourne in relation to environmental factors (G.S. Gardiner (1943a). Nevertheless, the correlative evidence West, 1909). Noteworthy of his style were accompanying obtained by Pearsall (1932) for seasonal silicate depletion photomicrographs of plankton communities in fields by diatoms in some English lakes was a major advance under low magnification (Fig. 2a). His untimely death in that has stood the test of time. In a wide-ranging review the influenza epidemic prevalent around 1919 was a major of freshwater algal ecology, Fritsch (1931) had raised the loss to the science. possibility of mass-recruitment of lake phytoplankton Contemporary with G.S. West was the early work from benthic stages. This feature was later demonstrated of F.E. Fritsch. Fritsch (Fig. 1d) had a training in classical in work on a partially planktonic (meroplanktonic) diatom botany in Germany, where this subject – including (Lund, 1954; Jewson et al., 1981), various cyanoprokaryotes freshwater algae – was far more advanced than in Britain. (Cyanobacteria, ‘blue-greens’: Preston et al., 1980; Head Most of his professional life was spent at the University et al., 1999), and a dinoflagellate (Heaney et al., 1983). of London, where he gained an unequalled reputation in ‘algology’ (later phycology). In 1903 he published a study Linkage to physiology and growth of the phytoplankton of the River Thames at London. dynamics Both he and G.S. West were impelled to consider the algae they observed in a geographical–environmental context, Evidence of natural factor-action complementary to as by the Wests for their favourite group, the desmids that obtainable from time-correlations is potentially (West & West, 1906b). Both took up time-sequences available from experimental physiology, provided that of algal occurrence as regular or irregular periodicity, extrapolation from laboratory to natural environment is the Wests dealing with the phytoplankton of some not extreme. Here the influence of W.H. Pearsall (Fig. 1c),

Fig. 1. Founders of freshwater phytoplankton ecology in Britain: (a) W. West; (b) G.S. West – from John et al. (2003); (c) W.H. Pearsall; (d) F.E. Fritsch; (e) J.W.G. Lund

DOI: 10.1608/FRJ-5.1.453 Freshwater Reviews (2012) 5, pp. 1-20 4 Talling, J.F.

Fig. 2. Approaches to the study of freshwater phytoplankton: (a) microscopy and floristic composition – photomicrograph by West & West (1906a) of net plankton from Lough Neagh; (b) quantitative sampling over longitudinal river sequences – the research vessel Malakal on the upper White Nile, 1953; (c) experimental manipulation – the ‘Lund tubes’ in Blelham Tarn, 1970. From Talling (2004). physiologist and ecologist based at Leeds University, nationally. Much physiology depended on the availability was historically decisive in conjunction with research of algal cultures, which the Chinese visitor Chu (1942) by J.W.G. Lund. Pearsall participated in following achieved for common freshwater phytoplankters in dilute growth increments over the vertical underwater light media, followed by more early application at Windermere gradient (Loose et al., 1934), and applied analysis of algal (Storey, 1944; Lund, 1949). growth in cultures (Pearsall & Loose, 1937) to advocate With respect to the logarithmic plotting of population the logarithmic interpretation of changes in planktonic increase or decrease, a British tradition was born, affecting populations (e.g. Pearsall et al., 1946). The latter was the interpretation of action by density-dependent and foreshadowed in a neglected paper of Scourfield (1897), density-independent factors. It was also represented but it was Pearsall’s influence that was responsible for in experiments to ‘bioassay’ the seasonal physical some major applications and extensions by Lund (1949, environment (light, temperature) for planktonic growth 1950) and subsequently by others at Windermere and by exposure of dilute, nutrient-replete cultures of

© Freshwater Biological Association 2012 DOI: 10.1608/ FRJ-5.1.453 Freshwater phytoplankton ecology 5 characteristic species exposed in situ over a depth-range origins influenced by Fritsch, Pearsall and the German (Lund, 1949; Talling, 1955; Cannon et al., 1961). scholar-refugee E.G. Pringsheim, and led to new Interpretation was helped by parallel exposure in the generations of interested students. Principal figures were laboratory (Talling, 1955). Later there were independent G.E. Fogg in London and later Menai Bridge in Wales, a experimental studies of growth dynamics of common one-time colleague and admirer of Fritsch and Pearsall, species, especially at Windermere (Hughes & Lund, who gave a wide attention to phytoplankton ecophysiology 1962; Jaworski et al., 1981, 2003; Butterwick et al., 1982, (Fogg & Thake, 1987; Fogg, 1991), and his former colleagues 2005), and by Gibson, Foy and associates near Lough and blue-green specialists W.D. P. Stewart at Dundee, A.E. Neagh in Northern Ireland (e.g. Foy, 1983; Gibson & Foy, Walsby at Bristol, and P. Fay – a Hungarian émigré – in 1988, 1989; Foy & Gibson, 1993; Richardson et al., 2000). London (Fig. 3). Teaching at London by another émigré – J. Nutrient inadequacy in natural waters was more Rzóska from Poland, a veteran of European hydrobiology frequently addressed throughout Britain from differences – generated interest in several subsequent British workers of algal growth in laboratory bioassays, using water on phytoplankton: A.E. Bailey-Watts, T.J. Lack and C.S. samples to which combinations of potentially limiting Reynolds. Also near London there was research on nutrients were added and the growth of indigenous or freshwater algae from Royal Holloway College by J.H. inoculated test algae followed. Both absolute and relative Evans and J.D. Dodge; at Bristol University, by F.E. Round (logarithmic) measures of growth were used, the former and his students, including B. Moss and C.M. Happey. as absolute increments of biomass or its correlate cell Later, eventually at Dundee University, G.E. Codd number, the latter as ‘cell divisions’ (= log2 increments or pursued the features of toxins produced by many doublings; e.g. Lund et al., 1971a, b). Dividing cells could blue-greens (Codd, 1984) and J.A. Raven mainly the be readily recognised in large dinoflagellates such as Ceratium hirundinella, and in one English lake – Esthwaite Water – their relative frequency over depth in a day-night cycle could be translated into a direct measure of population growth rate (Heller, 1977a; Frempong, 1982). After 1950 schools of algal physiology developed in several British universities and, directly or indirectly, applied their methods or findings to issues Fig. 3. The ‘Fogg school’ or Westfield College group of algal physiologists, with much input into of phytoplankton phytoplankton ecology, during a reunion at Dundee in 1976. From left: W.D.P. Stewart, G.E. Fogg, A.E. ecology. Many had Walsby and P. Fay. From Walsby (2006).

DOI: 10.1608/FRJ-5.1.453 Freshwater Reviews (2012) 5, pp. 1-20 6 Talling, J.F. subcellular aspects of algal growth, solute transport and by Steemann-Nielsen of a sensitive 14C method, and a photosynthesis (Raven & Geider, 1988). Physiological world-wide project to assess global aspects of organic work was also amplified or initiated at institutes by production – the International Biological Programme (IBP, Windermere (the Freshwater Biological Association, FBA) 1964–1974). Work on freshwater phytoplankton in Britain, and Lough Neagh (Department of Agriculture and its much affiliated to the IBP and using oxygen metabolism in successors, Northern Ireland; Freshwater Laboratory, productive waters, was carried out by Bindloss (1974, 1976) New University of Ulster). These could extend down to on Loch Leven, by Kowalczewski & Lack (1971) and Lack enzyme systems (e.g. phosphatases) involved in nutrient (1971) on the River Thames, and by Jewson (1976) and acquisition. Highly relevant work was also done at Jones (1977) on Lough Neagh in Northern Ireland. From some marine institutes, as by M.R. Droop at the Millport these some novel aspects emerged, including relation to

Marine Laboratory in Scotland (Tett et al., 1985). In the light climate and the pH–CO2 system and to topographic freshwater institutes, unlike the urban universities, it differentiation in the Thames and Lough Neagh. Other was operationally relatively easy to interrelate field and IBP work was made abroad under British auspices, laboratory approaches – as exemplified by studies at especially in tropical Africa (see later). Work from Windermere of Lund (1949), Talling (1957a, b, 1966a), various sites was extended by Mann (1975) and Morgan et Heaney et al. (1989) and Maberly et al. (2002), and at Lough al. (1980) to relations with secondary animal production. Neagh of Jewson et al. (1981) and Gibson & Foy (1988). In later years work involving planktonic photosynthesis and respiration was mainly associated with Light-relations: growth and issues of population dynamics (Heller, 1977b; Harris et al., photosynthesis 1979; Foy & Gibson, 1982; Reynolds et al., 1985) including buoyancy and sinking (Reynolds, 1975; Reynolds & Many studies on freshwater phytoplankton were Walsby, 1988), plus inhibitory effects of radiation near the influenced by earlier marine equivalents. This is especially water surface (Belay, 1981) and varying ability to take up true for photosynthetic performance and its relation to CO2 from depleted media (Talling, 1976; Saxby-Rouen light, first tested in British fresh waters with cultures of a et al., 1999; Maberly et al., 2009). The influence of marine diatom suspended in Loch Awe by Jenkin (1930). phytoplankton on light penetration underwater (Talling, Her work, during a summer vacation, was inspired by 1960; Jewson, 1977) was studied with improved spectral contact at Millport with the celebrated scientific duo S.M. resolution (Jewson et al., 1984). Physiological interests Marshall and A.P. Orr, whose similar studies in sea lochs were also extended to extracellular production of organic (Marshall & Orr, 1928) were remarkable pioneer work. solutes by Fogg and his associates (Fogg, 1971). His Her later marine work at Plymouth (Jenkin, 1937) was also work led internationally to diverse opinions on their outstanding for interrelating underwater photosynthesis quantitative magnitude and functional importance. with controlling energy fluxes, for which advice from the oceanographer W.R.G. Atkins was critical. It inspired Nutrient relations: natural and similar work with freshwater planktonic diatoms at cultured populations Windermere by Talling (1957a, b, 1966a), involving direct comparison of natural and cultured populations of Analyses of water for dissolved nutrients in low Asterionella formosa and depth-differentiation in the former. concentrations first became practicable in the 1920s, largely By this time photosynthesis had been widely from the attention of marine workers such as W.R.G. appreciated as a potential ‘rate-meter’ for assessing Atkins. They were applied in some following studies of organic production by planktonic communities. algal periodicity, as by Pearsall (1932) and Lind (1938). Applications were stimulated by the introduction in 1952 Pearsall established a correlation between diatom maxima

© Freshwater Biological Association 2012 DOI: 10.1608/ FRJ-5.1.453 Freshwater phytoplankton ecology 7 and severe silicate depletion in some English Lakes, later buoyancy and sinking behaviour (Lund, 1959) and, with confirmed by Mortimer (1939) and Storey (1944), and free-swimming by flagellates sensitive to factor-gradients, greatly extended by Lund (1950). Lund’s quantitative affected the reliability of population census. Their finer comparison of crop silica and available silicon (as SiO2) resolution could be obtained with novel sampling devices left little doubt that the latter was periodically amajor and photo-sensors (Heaney & Talling, 1980; Talling, limiting nutrient, but did not exclude the possibility of 1981); their understanding was aided by laboratory co-limiting factors. This circumstance was evident in experimentation (Rother & Fay, 1979; Heaney & Furnass, work by Gardiner (1943b) on diatom growth in London 1980; Fay, 1988; Heaney et al., 1989; Clegg et al., 2003). There reservoirs. Later work on diatom–silicon relationships and were conspicuous examples from buoyant blue-greens (e.g. budgets (Bailey-Watts, 1976; Gibson, 1981; Bailey-Watts et as ‘water-blooms’: Reynolds, 1973; Reynolds & Walsby, al., 1989) was aided by the restricted variation of Si in the 1988; Walsby, 1988), sinking diatoms (Heaney et al., biomass of most freshwater species and its relatively slow 1989; Davey & Heaney, 1989), and motile dinoflagellates recirculation. (George & Heaney, 1978; Heaney & Talling, 1980). Other major nutrient elements are nitrogen and Problems of multiple chemical forms, cellular storage phosphorus, applicable – unlike silicon – to all organisms and limnetic re-circulation complicated interpretations and potentially available in several chemical forms. The of the relation between soluble reactive phosphorus and latter have complicated comparisons and deductions phytoplankton periodicity. Here the oft-neglected work (e.g. by Lund, 1950, 1965) from nutrient budgets and the at Windermere of Mackereth (1953), on Asterionella formosa, relative amounts environmentally available and those was fundamental. He demonstrated capacity for variable incorporated in algal populations, of which different types P-storage in cells of both cultured and natural populations may have different preferences or absolute capacities. and the very low minimum ‘cell quota’ (Tett et al., 1985) N-fixation from molecular nitrogen had been established for growth. The exact significance of correlative field by De and Fogg for non-planktonic blue-greens, and later relationships between phytoplankton abundance and application to phytoplanktonic members were made ambient P concentrations was, in consequence, dubious. possible by use of 15N tracer or the acetylene-reduction This was reinforced by evidence of Rigler and associates assay. These applications were especially applied by Fogg in Canada for rapid circulation and exchange of so-called and Stewart and their associates (Stewart, 1969; Horne & ‘inorganic phosphate’ in lakes, hailed by some as a changed Fogg, 1970; Stewart & Alexander, 1971). Nevertheless the paradigm of thinking. Later correlations between total P capacity to utilise molecular nitrogen was not established concentration and phytoplankton centred on measures for some common planktonic blue-greens. Historically of total phytoplankton abundance (e.g. as chlorophyll a), the water-analytical information was better for low as originally proposed by Sakamoto and Vollenweider concentrations of nitrate- than of ammonium-nitrogen. In abroad and given numerous later applications in Britain most little-polluted British lakes the former was much the (e.g. Heaney et al., 1986, 1992); this did little to elucidate more abundant and received most attention. Its common the role of P-limitation in the ecology of individual species. summer depletion was influenced by denitrifying bacteria In the 1970s and later the issues of nutrient enrichment as well as by algae plus cyanoprokaryotes. Sedimentation (‘eutrophication’) of inland waters were prominent in was shown to be predominant in a correlative summer Britain, as elsewhere. Experimental bioassays of water relationship of the abundant Tychonema (formerly samples were often applied, based on the growth of test Oscillatoria) bourrellyi and low nitrate-nitrogen in organisms or of native species (e.g. Lund et al., 1971a, b). Windermere South Basin (Heaney et al., 1996). Unique in structure and length of operation were the Other distributions of phytoplankton in space, vertically very large experimental mesocosms (‘Lund tubes’: Fig. 2c) or horizontally, were of intrinsic interest in relation to established in Blelham Tarn, Cumbria, wherein physical

DOI: 10.1608/FRJ-5.1.453 Freshwater Reviews (2012) 5, pp. 1-20 8 Talling, J.F. and chemical manipulations (e.g. isolation from sediments, Windermere between the rise and fall of species-popula- vertical mixing, nutrient enrichment) could be applied (Lack tions of Rotifera and phytoplankton (Edmondson, 1965); & Lund, 1974; Lund & Reynolds, 1982; Reynolds, 1986). also from measures of gross community abundance based This work followed experiments with smaller cylinders and on net phyto- and zooplankton samples taken over lake partitions. Artificial de-stratification applied to the lake several decades and related to climatic changes induced by had radically altered the seasonal occurrence of a common Atlantic air- and water-circulation (George & Taylor, 1995). planktonic diatom prone to sedimentation (Lund, 1971b). Better quantitative measures of zooplankton One index of limitation, from the ratios (‘stoichiometry’) components, based on the partly published of nutrient elements – C, N and P – in particulate matter species-resolved counts of W.J.P. Smyly at Windermere, (seston) that included phytoplankton, and generalised as and of phytoplankton from long-term estimations there the Redfield ratio, was used occasionally (Tett et al., 1985; of chlorophyll a (Talling, 1993), enabled recognition in Heaney et al., 1986; Talling, 1993); it was liable to weighting two English Lakes of an early summer ‘clear-water phase’ by the characteristics of individual taxonomic groups. of reduced phytoplankton related to a corresponding Although it became widely accepted that low P input was maximum of parthenogenetic Daphnia of the hyalina-galeata the commonest agent of limitation, after the year 2000 this complex (George et al., 1990; Talling, 2002, 2003). In was increasingly challenged by evidence for N-limitation experimental enclosures (mesocosms) established in in some types of water-body (Maberly et al., 2002). another lake, more intensive short-term observation led The differences between species-occurrence in various to evidence of ‘tracking’ by zooplankton components types of fresh water, e.g. base-poor and base-rich, raised of specific phytoplankton components and to the the possibility that, apart from the classical ‘nutrients’, derivation of a ‘community grazing index’ of filtering major ions were significant chemical factors, either singly capacity (Thompson et al., 1982). However, much or in their ratios. Experimental work with cultures carbon flux to zooplankton could be independent of indicated that ratios per se were rarely important (Moss, phytoplankton, as shown by resolution using stable 1972), that K+ was but rarely likely to be a limiting nutrient isotopes in later work on Loch Ness (Jones et al., 1998). (Jaworski et al., 2003), and that correlated variables like From the 1940s to the end of the century the pioneer pH (Talling, 2011) and other CO2-system components work of H.M. Canter on chytrids and protozoans including free CO2 (Talling, 1976; Saxby-Rouen et parasitic on phytoplankters enabled this source of al., 1999) were locally or periodically significant. periodic and species-specific loss to be recognised in limnology (Canter & Lund, 1948; Canter, 1949, 1979; Grazing and parasitism Jewson et al., 1981). The incidence of algal viruses was recorded but little known (Daft et al., 1970). Quantitative issues of consumption (‘grazing’) of freshwater phytoplankton by micro-Crustacea, Rotifera Comparative site-extension and Protozoa were neglected in Britain for much of the 20th century. Its existence was known from qualitative Within Britain a diverse geology and topography led observations on gut contents and those of Pennington to water-bodies and drainage in varied mountainous (1940) on the collapse of a tub population (of ‘Diogenes’) to lowland basins that some pioneer limnologists linked to grazing by micro-Crustacea. Later observations in Europe classified respectively as Caledonian and from the English Lakes involved various Protozoa as Baltic lake types, divergent in their phytoplankton and consumers (Canter & Lund, 1968; Goulder, 1972; Finlay et environmental conditions. They differed, for example, in al., 1988; Canter et al., 1990). There were also indications the representation of desmids (West & West, 1906b; Lind from correlations deduced from long-term sampling on & Pearsall, 1945). Other early plankton surveys, in Ireland,

© Freshwater Biological Association 2012 DOI: 10.1608/ FRJ-5.1.453 Freshwater phytoplankton ecology 9 were of the large Lough Neagh by Dakin & Latarche was the possibility or actuality of transition from macro- (1913), of Lough Derg by Southern & Gardiner (1926), of phyte-dominated to plankton-dominated stable states other lakes by Lind & Pearsall (1945) and Round & Brook (Moss, 2003), of considerable theoretical and practical (1959). Pearsall (1923, 1932; Pearsall & Pearsall, 1925) significance. Until the 1980s the smallest fraction of plankton, explored differences linked to environment and season in the picoplankton of ≤2 µm cell size (Happey-Wood et al., the English (Cumbrian) Lakes, links that were expanded 1988), was generally overlooked, although it had been there by later studies reviewed in Macan (1970) and Talling recognised as a major component ‘of bacterial size’ in Loch (1999), some concentrating on an individual algal group, Leven (Bailey-Watts et al., 1968) and is now acknowledged such as the desmids (Canter & Lund, 1966; Lund, 1971a) as significant and universal in natural waters. or the diatom genus Tabellaria (Knudson, 1954). Reynolds A fundamental issue is the capacity for phytoplankton & Irish (2000) published an integrated and extensively to develop in running waters (Reynolds, 1988a). Such illustrated monograph on the diverse phytoplankton development in the Thames was early established by of Windermere. There was also attention to lakes in Fritsch (1903), and its quantitative production in that Scotland (Maulood & Boney, 1980; George & Jones, river and some tributaries was assessed in several later 1987; Maitland, 1994; Jones et al., 1998), especially Loch studies (Swale, 1964, 1969; Kowalczewski & Lack, Leven (Bailey-Watts, 1974, 1978) and in Ireland, especially 1971; Lack, 1971; Lack et al., 1978). Work here and Lough Neagh (Gibson et al., 1971; Gibson, 1981; Gibson on the Severn (Reynolds & Glaister, 1993) established & Fitzsimmons, 1982; Wood & Smith, 1993) and in Wales the significance of local and lateral stretches as foci of (Happey-Wood, 1975; Happey-Wood et al., 1988). Floristic phytoplankton increase. Well-developed phytoplankton differences in community composition were tested asa was studied in some east-flowing rivers, partly in quantitative guide to nutrient- and base-status (e.g. Brook, relation to the modification of river-input to the sea (LOIS 1965). project: Marker & Collett, 1997; Skidmore et al., 1998). A smaller scale of lakes, ponds and tarns was Studies by British workers in distant countries well-represented in works on phytoplankton and other introduced new factor-situations associated with algae in highland and lowland Britain, as in Scotland latitude and climate. In part these related to initiatives (Brook & Woodward, 1956), Cumbria (Belcher et al., in developing, often tropical, countries, in part to new 1966; Finlay & Maberly, 2000), Somerset (Happey & support to international projects. The considerable Moss, 1967), the Shropshire–Cheshire meres (Swale, British work on lake phytoplankton in tropical Africa 1968; Reynolds, 1976, 1979), and the Norfolk Broads provided examples of seasonality dominated by water (Moss & Balls, 1989; Moss, 2003). There were studies of flow (hydrology) or change in stratification (reviewed by periodicity in ponds and reservoirs near London (Evans, Talling, 1986), winter-less conditions , and opportunities 1988) and elsewhere (Youngman et al., 1976; Harper & for the development of river phytoplankton when – as Bullock, 1982; Cmiech et al., 1984). The stages of initial in the Nile, studied from the University College, later colonisation in a new northern reservoir were also University of Khartoum (Brook & Rzóska, 1954; Prowse recorded (Atkinson, 1978). In many smaller water-bodies & Talling, 1958; Hammerton, 1972; Talling et al., 2009) – a short residence time of water was a dominant factor there is a combination of great length, shallow gradients (Brook & Woodward, 1956; Reynolds & Lund, 1988). and local reservoir storage (Fig. 2b). Studies from G.S. Base status varied widely, affecting phytoplankton West (1907) onwards (Fish, 1957; Coulter, 1963; Talling, composition, and marl lakes with abundant benthic 1966b; Evans, 1997) showed extensive development macrophytes provided some instructive peculiarities in the very large African lakes. Photosynthesis by (Talling & Parker, 2003; Pentecost, 2009). In these shallow phytoplankton (Talling, 1965; Ganf, 1975) was often waters, and in the similarly shallow Norfolk Broads, there found to involve high rates per unit index of biomass

DOI: 10.1608/FRJ-5.1.453 Freshwater Reviews (2012) 5, pp. 1-20 10 Talling, J.F. and per unit area; reaction upon the water environment in basins of Windermere (40–45 years), of Reynolds & could be considerable. There were distinct assemblages Irish (2000) for changes in Windermere as a whole, and in chemically distinctive soda-lakes with a history of of Heaney et al. (1988) for changes in abundance of the evaporative loss (Jenkin, 1936; Wood & Talling, 1988) and dinoflagellate Ceratium spp. The lake Grasmere over 40 with distinctive nutrient regime (Wood et al., 1984), and in years has illustrated the consequences of changing sewage small upland waters (Lind, 1968). Between these extremes input and low but variable retention time (Reynolds and the relatively base-poor waters of Malaya (Prowse, & Lund, 1988; Reynolds et al., 2012). Elsewhere, long 1962) there were major differences in the phytoplankton. records from Loch Leven (Scotland) and Lough Neagh Other British work concerned the production of (Ireland) have been interpreted by Bailey-Watts (1978) and phytoplankton at very high, polar, latitudes, with limitations Gibson (1981; Gibson & Fitzsimmons, 1982) respectively. by low and very seasonal radiation input, accentuated by A distinct and much longer type of record is that snow and ice cover. Here the main British studies were on available from algal remains – especially of resistant small lakes on Signy Island by a team of the British Antarctic diatom frustules – in dated layers of lake sediments Survey; the work, also involving B. Heywood and J. Light, (palaeolimnology). In Britain late- and post-glacial has been reviewed by Priddle et al. (1986) and Fogg (1998). deposits are primarily involved. This was first developed at Windermere by Pennington (1943), who established Long-term records correlative relation between mass development of the diatom Asterionella formosa in Windermere and 19th Long-term sequences of sampling have been maintained century enrichment. She and later R.W. Battarbee – who by various individuals and organisations such as the had studied the sedimentary record of Lough Neagh Freshwater Biological Association. Examples are treated (Battarbee, 1977) – established active research groups at in a recent symposium (Maberly & Elliott, 2012). In this Windermere and London respectively. At the former respect the British record is exceptionally rich. Advantages Haworth (1980) was, among other work, able to relate are improved possibilities for correlating factors and the recent sedimentary record from Blelham Tarn with responses, wider factor-ranges, and the detection of an on-going long-term sampling record organised by responses to trends in climate (Thackeray et al., 2008; Nõges Lund. There were also studies from London of sites et al., 2010) and more especially nutrient enrichment. The overseas (Flower, 1994, 2001), and contributed sequences last, as eutrophication, has been a focus with influence from further British sites, including a loch in northern on the abundance and composition of phytoplankton. Scotland (Pennington et al., 1972), and work by McGowan Cumbrian lakes have provided several examples (Lund, et al. (1999) from Liverpool University on the Shropshire– 1978; Talling & Heaney, 1988). Cheshire meres that demonstrated a long historical The longest records are for the Cumbrian lakes, presence of blue-greens in lakes later noted for their blooms. especially Windermere, Esthwaite Water, Blelham Tarn and Grasmere. Counts of species have been followed in Models of phytoplankton production the first three since initiation by Lund in 1945, and total and relation to environmental abundance as chlorophyll a since 1964 (Talling, 1993). regime The latter used a 26-year sequence to summarise variants of gross seasonal abundance in four lake basins and Early recognition of regularities in depth-profiles of rates their relations with constituent species and particulate of photosynthesis, and relation to surface input of solar C, N and P. Example-deductions from the former are of radiation with near-exponential attenuation underwater, Lund (1978) for Blelham Tarn (32 years), of Neale et al. enabled the photosynthetic production per unit area (1991) and Maberly et al. (1994) for Asterionella formosa over longer time periods to be integrated and modelled

(Talling, 1957b, 1971). This approach was applied to (Spain) and Rigler (Canada). Similar divergence applied several British lakes during the International Biological to the extent of auto-induced community succession in Programme. An important variable was the mixed depth time (Reynolds, 1988b) and its frequency. Environmental of the water-column. In London reservoirs this could be changes classed as ‘intermediate disturbances’ have manipulated for practical ends, and Steel (1972) applied been considered as sources of taxonomic diversity, in further modelling to determine the consequences for the ‘intermediate disturbance hypothesis’ applied to phytoplankton growth. There were related actual results phytoplankton (Reynolds et al., 1993). from the experimental mesocosms in Blelham Tarn. In these and other more abstract generalisations, Aspects of the integration, as the model PROTECH, refutation of a hypothesis may be difficult or inherently were later applied by Reynolds and his associates impracticable. Others – most notably, and successfully, (reviewed in Reynolds et al., 2001) to specific growth Lund (e.g. 1949, 1965) – have preferred to concentrate rates over a corresponding depth-profile, and rates of on autecology linked to causal analysis. Another change of lake populations predicted using information approach, called ‘envelope ecology’ by Talling (1993), on the light climate, underwater light penetration, vertical has considered objective measures of total abundance of distribution of phytoplankton, species-specific growth mixed communities and explored the bases of possible characteristics, nutrient demands and estimates of specific regularities of periodic change, classified as ‘amplitude rates of loss processes (Reynolds et al., 1982) such as modulation’ and ‘frequency modulation’. Among these, respiration, sinking, grazing and mortality under nutrient the most intensely studied in Britain and elsewhere depletion. Differences of the variables between species – has been the spring diatom maximum, widespread in some probably related to size and morphotypes – could be temperate lakes and often dominated by a single species. shown to lead to seasonal preferences and species-succes- Other ‘British specialities’, discussed in previous sion, with greater sensitivity to some factors than to others. sections, have concerned: Comparison with the behaviour of a natural population • the logarithmic representation and interpretation of (‘validation’) could be close (Reynolds & Irish, 2000). population cycles; • long-term sequences of sampling; Higher-level generalisation • use of large experimental mesocosms; • culture growth-based assay in situ of physical factors; Over the past century those concerned with phytoplankton • episodes of microbial parasitism and determining science in Britain and elsewhere have differed in their factors; attitudes to useful generalisation. The classificatory • photosynthetic performance under diverse and tradition included division into trophic grades of lakes quantitatively characterised light-fields; (oligotrophic, mesotrophic, eutrophic) with correlations • spatial re-distribution of populations by buoyancy, to morphometry and plankton abundance. This was sedimentation, turbulence and active migration, their elaborated into finer lake typology by some Europeans laboratory replication and physiological regulation; but avoided by Pearsall. Other partially subjective • recruitment of some populations from resistant classifications have been applied to phytoplankton benthic stages – vegetative cells, cysts, akinetes; communities founded on taxonomic representation or • silicon budgets as insights on diatom ecology; recurrent associations of species (Reynolds, 1980). These • annual cycles, resources and productivity of tropical ‘higher level’ approaches and often compound entities were populations. variously considered (e.g. by Reynolds, Talling) as suited or Many early workers were impressed by dispersed unsuited for objective causal analysis – a difference that in plankton as a ‘model community’. This theme was later extremes has been represented internationally by Margalef taken up in Britain for phytoplankton, regarding it as

DOI: 10.1608/FRJ-5.1.453 Freshwater Reviews (2012) 5, pp. 1-20 12 Talling, J.F. a model photosynthetic cover (Talling, 1961, 1970), as of the eutrophic and non-stratifying Loch Leven (Kinross, a model of community succession at the cellular level Scotland). Journal of Ecology 66, 741-771. (Reynolds, 1997, 2006), and as a more general model for Bailey-Watts, A.E., Bindloss, M.E. & Belcher, J.H. (1968). population dynamics related to environmental factors Freshwater primary production by a blue-green alga of bacterial (Talling, 2002). In these respects its suitability is enhanced size. Nature 220, 1344-1345. by the possibilities of quantitative sampling, species Bailey-Watts, A.E., Smith, I.R. & Kirika, A. (1989). The dynamics of attribution and manipulation in cell suspensions of silica in a shallow, diatom-rich Scottish loch. II. The influence of populations that are generally more circumscribed than in diatoms on an annual budget. Diatom Research 4, 191-205. the marine environment; derivation of cultured material Battarbee, R.W. (1977). Observations of the recent history of and experimental approaches with natural and cultured Lough Neagh and its drainage basin. Philosophical Transactions populations; the prevalence of comparatively regular of the Royal Society (B) 281, 303-345. environmental gradients of light, temperature and chemical Belay, A. (1981). An experimental investigation of phytoplankton variables; a close involvement of biogeochemistry; and photosynthesis at lake surfaces. New Phytologist 89, 61-74. the frequency of population cycles over several orders of Belcher, J.H., Swale, E.M.F. & Heron, J. (1966). Ecological and magnitude in a short time-scale that have a basis in cellular morphological observations on a population of Cyclotella division, growth rate modulation and (Jewson et al., 1981; pseudostelligera Hustedt. Journal of Ecology 54, 335-340. Reynolds et al., 1982) quantifiable loss-processes. This is an Bindloss, M.E. (1974). Primary production of phytoplankton in array of advantages probably unequalled in present-day Loch Leven, Kinross. Proceedings of the Royal Society of Edinburgh quantitative ecology and population dynamics. 74, 157-181. Bindloss, M.E. (1976). The light climate of Loch Leven, a Acknowledgements shallow Scottish lake, in relation to primary production by phytoplankton. Freshwater Biology 6, 501-518. I am indebted to Chris Gibson, Ivan Heaney, Colin Brook, A.J. (1965). Planktonic algae as indicators of lake types, Reynolds and two anonymous referees for perceptive with special reference to the Desmidiaceae. Limnology and comments and suggestions. Photographic illustrations Oceanography 10, 401-411. were made available by the kindness of David John and Brook, A.J. & Rzóska, J. (1954). The influence of the Gebel Auliya The Phycologist (Fig. 1a, b), the Freshwater Biological dam on the development of Nile plankton. Journal of Animal Association (Fig. 1c, d, e; Fig. 2), and Tony Walsby (Fig. 3). Ecology 23, 101-114. Brook, A.J. & Woodward, W.B. (1956). Some observations on the References effects of water inflow and outflow on the plankton of small lakes. Journal of Animal Ecology 25, 22-35. Atkinson, K.M. (1978). The initial development of phytoplankton Butterwick, C., Talling, J.F. & Heaney, S.I. (1982). A comparison in Cow Green Reservoir (Upper Teesdale), a new impoundment of eight methods for estimating the biomass and growth of in Northern England. In: Algae and the Aquatic Environment (ed. planktonic algae. British Phycological Journal 17, 69-79. F.E. Round), pp. 30-43. Biopress, Bristol. Butterwick, C., Heaney, S.I. & Talling, J.F. (2005). Diversity in the Bailey-Watts, A.E. (1974). The algal plankton of Loch Leven, influence of temperature on the growth rates of freshwater Kinross. Proceedings of the Royal Society of Edinburgh (B) 74, algae, and its ecological relevance. Freshwater Biology 50, 135-156. 291-300. Bailey-Watts, A.E. (1976). Planktonic diatoms and silica in Loch Cannon, D., Lund, J.W.G. & Sieminska, J. (1961). The growth of Leven, Kinross: a one-month silica budget. Freshwater Biology Tabellaria flocculosa (Roth) Kűtz. var. flocculosa (Roth) Knuds. 6, 203-213. under natural conditions of light and temperature. Journal of Bailey-Watts, A.E. (1978). A nine-year study of the phytoplankton Ecology 49, 277-287.

Canter, H.M. (1949). The importance of fungal parasitism in and biological factors. Journal of Plankton Research 11, 1185-1199. limnology. Verhandlungen der Internationalen Vereinigung fűr Edmondson, W.H. (1965). Reproductive rates of planktonic theoretische und angewandte Limnologie 10, 107. rotifers as related to food and temperature in nature. Ecological Canter, H.M. (1979). Fungal and protozoan parasites and their Monographs 35, 61-111. importance in the ecology of the phytoplankton. Report of the Evans, J.H. (1988). Long term changes in planktonic Freshwater Biological Association 47, 43-50. Cryptophycean populations. In: Algae and the Aquatic Canter, H.M. & Lund, J.W.G. (1948). Studies on plankton parasites. Environment (ed. F.E. Round), pp. 44-52. Biopress, Bristol. I. Fluctuations in the numbers of Asterionella formosa Hassall in Evans, J.H. (1997). Spatial and seasonal distribution of relation to fungal epidemics. New Phytologist 47, 238-261. phytoplankton in an African Rift Valley lake (Lake Albert, Canter, H.M. & Lund, J.W.G. (1966). The periodicity of planktonic Uganda-Zaїre). Hydrobiologia 354, 1-16. desmids in Windermere. Verhandlungen der Internationalen Fay, P. (1988). Viability of akinetes of the planktonic Vereinigung fűr theoretische und angewandte Limnologie 15, 163-172. cyanobacterium Anabaena circinalis. Proceedings of the Royal Canter, H.M. & Lund, J.W.G. (1968). The importance of Protozoa Society (B) 234, 283-301. in controlling the abundance of planktonic algae in lakes. Finlay, B.F. & Maberly, S.C. (2000). Microbial diversity in Priest Pot: a Proceedings of the Linnaean Society of London 179, 203-219. Productive Pond in the English Lake District. Freshwater Biological Canter, H.M., Heaney, S.I. & Lund, J.W.G. (1990). The ecological Association, Ambleside. 73 pp. significance of grazing on planktonic populations of Finlay, B.F., Berninger, U.G., Clarke, K.J., Cowling, A.J., Hindle, cyanobacteria by the ciliate Nassula. New Phytologist 114, R.M. & Rogerson, A. (1988). On the abundance and distribution 247-263. of protozoa and their food in a productive freshwater pond. Chu, S.P. (1942). The influence of the mineral composition of the European Journal of Protistology 23, 205-217. medium on the growth of planktonic algae. Part I. Methods Fish, G.R. (1957). A seiche movement and its effect on the and culture media. Journal of Ecology 30, 284-325. hydrology of Lake Victoria. Fisheries Publication, Colonial Office, Clegg, M.R., Maberly, S.C. & Jones, R.I. (2003). Behavioural London 10, 1-68. responses of freshwater phytoplankton flagellates to a Flower, R. (1994). A review of current and recent environmental temperature gradient. European Journal of Phycology 38, 195-203. research on Lake Baikal from a British perspective. Freshwater Cmiech, M.A., Reynolds, C.S. & Leedale, G.F. (1984). Seasonal Forum 1, 8-22. periodicity, heterocyst differentiation and sporulation of Flower, R. J. (ed.) (2001). North African lakes. Aquatic Ecology 35, planktonic Cyanophyceae in a shallow lake, with special 259-484. reference to Anabaena solitaria. British Phycological Journal 19, Fogg, G.E. (1971). Extracellular products of algae in freshwater. 245-257. Archiv fűr Hydrobiologie Beihefte, Ergebnisse der Limnologie 5, 1-25. Codd, G.K. (1984). Toxins of freshwater cyanobacteria. Fogg, G.E. (1991). The phytoplanktonic ways of life. New Microbiological Sciences 1, 48-52. Phytologist 118, 191-232. Coulter, G.W. (1963). Hydrological changes in relation to Fogg, G.E. (1998). The Biology of Polar Habitats. Oxford University biological production in southern Lake Tanganyika. Limnology Press, Oxford. & Oceanography 8, 463-477. Fogg, G.E. & Thake, B. (1987). Algal Cultures and Phytoplankton Daft, M.J., Begg, J. & Stewart, W.D.P. (1970). A series of blue-green Ecology. 3rd edition. University of Wisconsin Press, Madison. algae from freshwater habitats in Scotland. New Phytologist 69, Foy, R.H. (1983). Interaction of temperature and light on the 1029-1038. growth rates of two planktonic Oscillatoria species under a short Dakin, W. & Latarche, M. (1913). The plankton of Lough Neagh. photoperiod regime. British Phycological Journal 18, 267-273. Proceedings of the Royal Irish Academy (B) 30 (3), 20-96. Foy, R.H. & Gibson, C.E. (1982). Photosynthetic characteristics of Davey, M.C. & Heaney, S.I. (1989). The control of sub-surface blue-green algae; the response of twenty strains grown under maxima of diatoms in a stratified lake by physical, chemical high and low light. British Phycological Journal 17, 189-193.

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Foy, R.H. & Gibson, C.E. (1993). The influence of irradiance, morphology of planktonic blue-green algae in an unstratified photoperiod and temperature on the growth kinetics of three lake (Lough Neagh, Northern Ireland). Internationale Revue der planktonic diatoms. European Journal of Phycology 28, 203-212. gesamten Hydrobiologie 66, 641-664. Frempong, E. (1982). The space-time resolution of phased cell Gibson, C.E. & Foy, R. H. (1988). The significance of growth rate division in natural populations of the dinoflagellate Ceratium and storage products for the ecology of Melosira italica ssp. hirundinella. Internationale Revue der gesamten Hydrobiologie und subarctica in Lough Neagh. In: Algae and the Aquatic Environment Hydrographie 67, 323-339. (ed. F.E. Round), pp. 88-106. Biopress, Bristol. Fritsch, F.E. (1903). Further observations on the phytoplankton of Gibson, C.E. & Foy, R.H. (1989). On temperature-independent the River Thames. Annals of Botany 17, 631-646. growth of phytoplankton. Journal of Plankton Research 11, Fritsch, F.E. (1931). Some aspects of the ecology of freshwater 605-607. algae (with special reference to static waters). Journal of Ecology Gibson, C.E., Wood, R.B., Dickson, E.L. & Jewson, D.H. (1971). 19, 233-272. The succession of phytoplankton in Lough Neagh, 1968–70. Fritsch, F.E. & Rich, F. (1913). Studies on the occurrence and Mitteilungen der Internationalen Vereinigung fűr theoretische und reproduction of British freshwater algae in nature. 3. A 4 years’ angewandte Limnologie 19, 146-160. observation of a freshwater pond. Annales de Biologie lacustre 6, Goulder, R. (1972). Grazing by the ciliated protozoan Loxodes 1-83. magnus on the alga Scenedesmus in a eutrophic pond. Oikos 23, Ganf, G.G. (1975). Photosynthetic production and irradiance- 109-115. photosynthesis relationships of the phytoplankton from a Griffiths, B.M. (1923). The phytoplankton of bodies of freshwater, shallow equatorial lake (Lake George, Uganda). Oecologia 18, and the factors determining its occurrence and composition. 165-183. Journal of Ecology 11, 184-213. Gardiner, A.C. (1943a). Measurement of phytoplankton Griffiths, B.M. (1939). Early references to waterbloom in British population by the pigment extraction method. Journal of the lakes. Journal of the Linnaean Society of London 151, 12-19. Marine Biological Association 25, 739-744. Hammerton, D. (1972). The Nile River – a case history. In: Gardiner, A.C. (1943b). Silicon and phosphorus as factors limiting River Ecology and Man (eds R.T. Oglesby, C.A. Carlson & J.A. development of diatoms. Journal of the Society of Chemical McCann), pp. 171-214. Academic Press, New York & London. Industry of London 60, 73-78. Happey-Wood, C.M. (1975). Distinctions in algal ecology and George, D.G. & Heaney, S.I. (1978). Factors influencing the spatial production in two linked upland lakes, Gwynedd, N. Wales. distribution of phytoplankton in a small productive lake. Verhandlungen der Internationalen Vereinigung fűr theoretische und Journal of Ecology 66, 133-155. angewandte Limnologie 19, 1045-1056. George, D.G. & Jones, D.H. (1987). Catchment effects on the Happey, C.M. & Moss, B. (1967). Some aspects of the biology of horizontal distribution of phytoplankton in five of Scotland’s Chrysococcus diaphanus in Abbot’s Pond, Somerset. British largest freshwater lochs. Journal of Ecology 75, 43-59. Phycological Bulletin 3, 269-279. George, D.G. & Taylor, A.H. (1995). UK lake plankton and the Happey-Wood, C.M., Kennaway, G.M.A., Ong, M.H., Chittenden, Gulf Stream. Nature, London 378, 139. A.M. & Edwards, G. (1988). Contributions of nano- and George, D.G., Hewitt, D.P., Lund, J.W.G. & Smyly, W.D.P. (1990). pico-algae to the productivity of phytoplankton and epipelic The relative effects of enrichment and climate change on the algae in an upland Welsh lake. In: Algae and the Aquatic long-term dynamics of Daphnia in Esthwaite Water, Cumbria. Environment (ed. F.E. Round), pp. 168-184. Biopress, Bristol. Freshwater Biology 23, 55-70. Harper, D.M. & Bullock, J.A. ed. (1982). Rutland Water – decade of Gibson, C.E. (1981). Silica budgets and the ecology of planktonic change. Junk, The Hague. diatoms in an unstratified lake (Lough Neagh, N. Ireland). Harris, G.P. (1986). Phytoplankton Ecology: Structure, Function and Internationale Revue der gesamten Hydrobiologie 65, 49-84. Fluctuation. Chapman and Hall, London. Gibson, C.E. & Fitzsimons, A.G. (1982). Periodicity and Harris, G.P., Heaney, S.I. & Talling, J.F. (1979). Physiological and

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