Phytoplankton production and growth rate in Lake Tanganyika: evidence of a decline in primary productivity in recent decades

1. This study focused on phytoplankton production in Lake Tanganyika. We provide new estimates of daily and annual primary production, as well as growth rates of phytoplankton, and we compare them with values published in former studies.
2. Chlorophyll- a (chl- a ) in the mixed layer ranged from 5 to 120 mg chl- a  m⁻² and varied significantly between rainy and dry seasons. Particulate organic carbon concentrations were significantly higher in the south basin (with 196 and 166 mg C m⁻³ in the dry and the rainy season, respectively) than in the north basin (112 and 109 mg C m⁻³, respectively).
3. Carbon : phosphorus (C : P) ratios varied according to season. Phosphorus limitation seemed to occur more frequently than nitrogen limitation, especially during the rainy season. Severe P deficiencies were rare.
4. Measured particulate daily primary production ranged from 110 to 1410 mg C m⁻² day⁻¹; seasonal contrasts were well marked in the north basin, but less in the south basin, where primary production peaks occurred also in the rainy season. Estimates of annual primary production, based on daily primary production calculated from chl- a and water transparency, gave values lower than those reported in previous studies. Picophytoplankton accounted on average for 56% of total particulate production in the south basin during the wet season of 2003.
5. Phytoplankton growth rates, calculated from primary production, ranged from 0.055 to 0.282 day⁻¹; these are lower than previously published values for Lake Tanganyika.     

The life-cycle and productivity of Tibifex tubifex (Oligochaeta; Tibificidae) in the Moat-Feeder Stream, Cardiff, South Wales

A population of Tubifex tubifex in an organically rich stream was found to have an annual life-cycle with a prolonged period of reproductive activity throughout the winter and spring. Cocoons were produced mainly during the late winter and early spring. No cocoons were found during August and September, and there were few mature worms at this time.
The population density ranged between 5420 m⁻² in mid-September and 613000 m ⁻² in mid-May. The maximum population biomass (Bmax) recorded was 106 g dry wt m⁻² (March) and the minimum was 10 g dry wt m⁻² (September). Total annual production (P) was 139 g dry wt m⁻² and the average annual biomass ( B ) was 46 g dry wt m⁻² giving an annual P/ B ratio of 3.0, and a P/Bmax ratio of 1.3.     

The effect of an emergent macrophyte (Typha angustifolia) on sediment resuspension in a shallow north temperate lake

1. The effects of emergent macrophytes on water turbidity and sediment resuspension in the shallow Kirkkojärvi basin of Lake Hiidenvesi were studied with sediment traps, and concomitant sediment and water samples. The study was conducted during May–August in three different zones of a stand of the emergent Typha angustifolia .
2. Within the stand (5 m from the edge), both the concentration of suspended solids and the rate of sediment resuspension were significantly lower than at the edge and outside the stand (5 m from the edge). The differences between the zones increased towards the end of summer together with the growing stem density. During the study period (82 days), 2210 g dw m⁻² of sediment was resuspended in the outer zone. At the edge and in the inner zone, the corresponding numbers were 1414 and 858 g dw m⁻², respectively.
3. With the resuspended sediment, 39.4 mg P  m⁻² day⁻¹ was brought to the water column outside the stand, 22.4 mg P  m⁻² day⁻¹ at the edge and 13.4 mg P  m⁻² day⁻¹ within the stand.
4. In early summer, the concentration of suspended solids had a highly significant positive effect on soluble reactive phosphorus (SRP) concentration in the water, whereas in late summer no effect was found. During the study period, phosphorus retention by emergent macrophyte stands corresponded to 3–5% of the present annual external phosphorus loading of the Kirkkojärvi basin.     

Productivity of a herbivorous pupfish population (Cyprinodon nevadensis) in a warm desert stream

Production and food intake by an herbivorous pupfish population ( Cyprinodon nevadensis amargosae ) living in the outflow of a thermal artesian well (Tecopa Bore) near Death Valley, California is described. Water issues from the ground at 47.5° C and cools 8.12° C before leaving the study area 300 m from the source. High stream temperatures restricted the pupfish population to some 41 % of the study area, with a resulting mean density of 89 individuals m⁻² (range = 13 to 196m⁻²). Biomass estimates ranged from 7 kcal m⁻² to 42 kcal m⁻². The mean annual standing crop of pupfishes (24 kcal m⁻²) turned over about five times annually. Growth rates were highest in juveniles (♂= 9.5% day⁻¹) and slowest in large adults (♀= 08% day⁻¹). Monthly production ranged from 22 kcal m⁻² in September to 3 kcal m⁻² in July and August. Pupfish in Tecopa Bore fed on algae and detritus, ingesting 1941 kcal m⁻² yr⁻¹ or 17.5% of the annual net primary production. 119 kcal m⁻² yr⁻¹ was deposited in growth. This latter value is approximately ten times greater than values previously reported for large carnivorous fishes but is comparable lo values reported for herbivorous fishes under pond culture.     

Very high secondary production at a lake outlet

SUMMARY. 1. Larvae and pupae of Simulium noelleri Fried, coated the concrete of parts of an artificial lake outlet in southern England.
2. In the first two (of three) summer generations, development was synchronous and this allowed the calculation of their secondary production by the instantaneous growth method. The production of the two summer generations was, respectively, 229.1 g C m⁻² (7.4 g C m⁻²day⁻¹) and 185.5 g C m⁻² (8.8 g C m⁻² day⁻¹) The contribution of the third summer generation, and the overwintering generation, to annual production would be less than that of the first two summer generations. Nevertheless, annual production will have exceeded 500 g C m⁻² at this site.
3. Larvae are suspension feeders and they captured the rich supply of particulate and dissolved organic material which passed over them after export from the lake. As food is brought to the larvae they only require space for attachment and can thus build up very high population densities (which exceeded 1 × 10⁶ m⁻² on some occasions during the summer). The high population densities result in a high biomass and hence in the high levels of production.     

Seasonal changes in vertical distribution, biomass and faecal production of tubificids in the profundal region of a shallow Japanese lake

Two tubificid species Limnodrilus hoffmeisteri and L. claparedeianus formed more than 93% of the total number of oligochaetes in the profundal. Limnodrilus spp. worms were found down to 33 cm in the sediment but in great numbers in the upper zone in June and October. Worms confined to the top 15 cm of sediment accounted for 53-92% of the total number. There were two annual maxima in population density and biomass, one in late spring (66000 inds m⁻², 17 g wet wt m⁻²) and the other in mid autumn (97000 inds m⁻², 176 g wet wt m⁻²). Two regression lines describing the effect of temperature on faecal production rate were obtained; Log F = 0.0604 T (°C) −0.7660 (below 15°C), Log F = 0.0266 T – 0.2170 (above 15°C). In total 26.8 kg dry wt m⁻² of sediment was defecated annually by Limnodrilus spp. The sediment in the 0–10 cm stratum may pass through the guts of the worms 2.3 times a year. Sedimentation rates in profundal region were very low with respect to the faecal production rates of the tubificids.     

DEMOGRAPHY OF FRONDS OF SARGASSUM LAPAZEANUM FROM SOUTHERN BAJA CALIFORNIA, MEXICO, DURING EARLY STAGES

In terms of biomass, Sargassum lapazeanum (Phaeophyceae, Fucales) is one of the most important seaweeds in La Paz Bay, on the southeastern coast of the Baja California Peninsula, Mexico. This species can be found year-round in shallow subtidal sites. Standing biomass is lowest between fall and winter and highest between spring and summer. We are currently studying the annual demography of fronds as a necessary step to understanding the mechanisms of population regulation. Work is being done on a bed of about 90 m in length and three m in width. At this stage of abstract submission, we can report results for winter only. Random samples were collected in February and in March 2000 to estimate stand biomass, frond density, and size (frond length) structure. In addition, we labeled fronds with numbered plastic tags to estimate their rates of growth and of mortality during this period. Rates of recruitment were estimated from a combined analysis of the above. In February, fronds were all shorter than 6.5 cm. Between February and March, mean total frond density increased from 122 to 776 fronds m⁻² (n = 30 quadrats). The mean recruitment rate was 667 fronds m⁻², the mean growth rate was 0.5 cm day⁻¹ (n = 60 fronds), and the mean mortality rate was 43 fronds m⁻². From last year's preliminary observations, we expect to observe peaks of biomass and of reproduction in late spring, followed by negative growth rates and high mortality rates during summer.     

Production and litter processing by crayfish in an Appalachian mountain stream

SUMMARY 1. Mean annual density and biomass () of Cambarus bartonii in an Appalachian mountain stream (U.S.A.) was 12 individuals m⁻² and 1669 mg (ash-free dry weight) m⁻².
2. Annual production ( P ) of C bartonii was 961 mg AFDW m⁻². Despite high biomass, low growth rates resulted in low production and a low P/ ratio of 0.58.
3. While C bartonii constituted 61% of the total macroinvertebrate biomass, it contributed only 13% of annual community secondary production.
4. Litter processing was positively related to temperature and crayfish size. Cambariis bartotnii was estimated to comminute 36 g m⁻² y⁻¹ of leaf litter (>1 mm²) to 24 g m−^>2 y⁻¹ fine particulate material (<1 mm²). The annual pattern of litter comminution by crayfish was regulated by temperature. As a result, >5()% of shredding activity by crayfish occurred from June to September which was also the period of lowest litter standing crops and activity of other shredding macroinvertebrates.
5. We speculate that during summer crayfish play an important role in temperate woodland streams by converting slowly processed leaf litter species (e.g. Rhododendron ) to fine particles which are then available to collector-gatherers (e.g. Chironomidae, Oligochaeta).     

Phytoplankton ecology in an Antarctic lake

SUMMARY. The ecology of the phytoplankton of Heywood Lake, Signy Island, South Orkney Islands, Antarctica was investigated during 1969–72. The lake, which is ice-covered for 8–10 months per year, is moderately eutrophic due to enrichment by seal excreta.
The annual cycle of the phytoplankton is described. During the winter (approximately May-September), very few algal cells could be detected in the water column and ¹⁴C fixation was below measurable limits. In spring (October-November), a rapidly-growing population of algae caused a large increase in the chlorophyll- a concentration (maximum value 170 mg m⁻²) but carbon fixation remained low, with values <500 mg C m⁻² day⁻¹. The algae contributing to this peak were mainly small chlorophytes and chrysophytes. The summer open-water period (December-March) was characterized by a different phytoplankton population dominated by cryptophytes. Chlorophyll levels were lower ( c . 40 mg m⁻²) but ¹⁴C fixation rates >3 g C m⁻² day⁻¹ were measured on bright days. Values for Assimilation Number were very high (maximum value 10.5 mg C h⁻¹ mg⁻¹ (chlorophyll- a ) in January (1971) though temperatures never exceeded 8°C. In autumn, the phytoplankton regressed to winter levels. Both spring and summer algal populations probably overwinter as resting stages.     

Modelling black fly production dynamics in blackwater streams

SUMMARY. 1. Two predictive models were employed along with intensive field sampling to estimate production of black flies ( Simulium spp.) on snags (submerged wood) in three blackwater streams on the Georgia Coastal Plain of the southeastern U.S.A. One model predicts daily growth rate from temperature and hydrograph pattern; the other predicts habitat abundance (of snags) from river height.
2. In the sixth order Ogeechee River, annual production was twice as high in 1982 (7.1 g dry mass [=DM] m⁻² of snag surface) as in 1983 (3.6 g DM m⁻²). When converted to production per m² of river bottom, values were 35–40% of the snag surface estimates. Annual production was much lower in fourth order Black Creek (1982, 1.3 g DM m⁻² of snag surface) and much higher in the sixth order Satilla River (1975, 15.6–40.0 g DM m⁻²).
3. There was a distinct bimodal pattern of black fly production in the Ogeechee River in both years, with peaks occurring in winter and summer. Similar bimodal patterns of production were found in Black Creek and in the Satilla River. Although there appears to be an intrinsic component to the bimodal pattern, production peaks (growth rate and biomass) appear to be associated with initial stages of flooding.
4. Annual production/biomass ratios (37–85) are the highest reported for black fly populations. The variation of annual P/B ratios among sites was more strongly dependent on the temporal distribution of standing stock biomass than on differences in growth rates. Variation in production among sites appears to be due to differences in current velocity, hydro-graph variability, and abundance of coexisting consumers.     

Development of leaf thickness for Plectranthus parviflorus – Influence of photosynthetically active radiation

Photosynthetically active radiation (PhAR) is apparently the environmental factor having the greatest influence on leaf thickness for Plectranthus parviflorus Henckel (Labiatae). A four-fold increase in leaf thickness from 280 to 1170 μm occurred as the PhAR was raised from 1.3 to 32.5 mol m⁻² day⁻¹. Compared to a constant PhAR of 2.5 mol m⁻² day⁻¹, a PhAR of 32.5 mol m⁻² day⁻¹ for one week during the first week (with return to 2.5 mol m⁻² day⁻¹ during the second and third weeks) led to an increase in final leaf thickness by 323 μm (to 802 μm). When increased PhAR was applied during the second week the increase in final thickness over the control was 217 μm, and when increased PhAR was applied during the third week it was 99 μm. However, leaf thickness was not simply responding to total daily PhAR, since a leaf 450 μm thick could occur at a low instantaneous PhAR for a long daytime (total daily PhAR of 1.5 mol m⁻² day⁻¹) and at a high PhAR for a short daytime (4.5 mol m⁻² day⁻¹). Total daily CO₂ uptake (net photosynthesis) was approximately the same in the two cases, suggesting that this is an important factor underlying the differences in leaf thickness. Leaf thickness is physiologically important, since thicker leaves tend to have greater mesophyll surface area per unit leaf area ( A ^mes/ A ) and hence higher photosynthetic rates.     

Population dynamics and production of Jesogammarus annandalei, an endemic amphipod, in Lake Biwa, Japan

SUMMARY 1. Population dynamics and production of Jesogammarus annandalei , an endemic amphipod in Lake Biwa, were examined from April 1997 to June 1998. The life cycle of this species was 1 year with the new generation beginning in early autumn. They preferred low temperature (<12 °C) and their spatial distribution varied seasonally and accordingly.
2. In deep water, the abundance of J. annandalei ranged from 200 to 63 000 m⁻² and decreased towards summer and the biomass (0.01∼3.6 g C m⁻²) was on average comparable that of zooplankton. The density was much higher than that recorded by a study conducted 35 years ago.
3. Individual growth rate of this amphipod was high in winter and spring but decreased in summer. Annual production of J. annandalei (6.2 g C m⁻² year⁻¹) was only 2% of primary production but was at the higher end of the range reported for amphipods in oligo- and mesotrophic lakes.
4. These results are consistent with the view that Lake Biwa is becoming more eutrophic, with a consequent decrease in the abundance of predatory fish in the profundal zone.     

A study of the dynamics of a natural population of the freshwater crayfish, Austropotamobius pallipes

SUMMARY. 1. Population size, survival and recruitment were estimated from mark—recapture data at frequent intervals.
2. Two types of mortality were identified: (1) moult-associated—an intense but short duration mortality, and (2) overwinter mortality, which operated at a lower estimated weekly rate but over some 30 weeks.
3. Population size reached a mid-summer peak as recruitment into the catchable size classes occurred. Thereafter numbers declined slowly to the end of the growth season, followed by an overwinter decline of some 50% of total.
4. Annual production was estimated at 196.5 kg wet wt ha⁻¹ in 1977, 116.8 kg wet wt ha⁻¹ in 1978 and 87.79 kg wet wt ha⁻¹ in 1979.     

A simple model of the eco-hydrodynamics of the epilimnion of Lake Tanganyika

1. The ecosystem response of Lake Tanganyika was studied using a four-component, nutrient–phytoplankton–zooplankton–detritus, phosphorus-based ecosystem model coupled to a nonlinear, reduced-gravity, circulation model. The ecosystem model, an improved version of the earlier eco-hydrodynamics model developed for Lake Tanganyika, was used to estimate the annual primary production of Lake Tanganyika and its spatial and temporal variability. The simulations were driven with the National Centres for Environmental Protection (NCEP) records for winds and solar radiation forcing.
2. The simulated annual cycles of the four ecosystem variables and the daily net primary production were compared with the observations. The comparison showed that simulations reproduced realistically the general features of the annual cycles of epilimnial phosphate, net primary production and plankton dynamics.
3. The climatic simulations for the years 1970–2006 yielded a daily averaged integrated upper layer net production ranging from 0.11 to 1.78 g C m⁻² day⁻¹ and daily averaged chlorophyll- a (chl- a ) from 0.16 to 4.3 mg m⁻³. Although the nutrient concentrations in the epilimnion during the strong wind years were high, the net production was low, which is partly because of the greater vertical mixing, produced by strong winds, exposing the phytoplankton to low light conditions in deeper waters. The simulated annual net production and chl- a agreed quite well with observed production available in the literature.
4. We envisage using this model to predict the future scenarios of primary productivity in the lake.     

Temporal dynamics in epipelic, pelagic and epiphytic algal production in a clear and a turbid shallow lake

SUMMARY 1. Pelagic and epipelic microalgal production were measured over a year in a pre-defined area (depth 0.5 m) in each of two lakes, one turbid and one with clear water. Further estimates of epiphytic production within reed stands were obtained by measuring production of periphyton developed on artificial substrata.
2. Total annual production of phytoplankton and epipelon was 34% greater in the turbid lake (190 g C m⁻² year⁻¹) than in the clearwater lake (141 g C m⁻² year⁻¹). However, the ratio of total production to mean water column TP concentration was two fold greater in the clearwater lake.
3. Phytoplankton accounted for the majority of the annual production (96%) in the turbid lake, while epipelic microalgal production dominated (77%) in the clear lake. The relative contribution of epipelic algae varied over the year, however, and in the turbid lake was higher in winter (11–25%), when the water was relatively clear, than during summer (0.7–1.7%), when the water was more turbid. In the clearwater lake, the relative contribution of epipelon was high both in winter, when the water was most clear, and in mid-summer, when phytoplankton production was constrained either by nutrients or grazing.
4. Compared with pelagic and epipelic primary production, epiphytic production within a reed stand was low and did not vary significantly between the lakes.
5. The study supports the theory of a competitive and compensatory trade-off between primary producers in lakes with contrasting nutrient concentrations, resulting in relatively small differences in overall production between clear and turbid lakes when integrating over the season and over different habitats.     

Population growth rates of two closely related rotifer species: effects of food quantity, particle size, and nutritional quality

SUMMARY. 1. The relationship between population growth rates and the concentrations of several algal species was determined in laboratory experiments with the rotifers Brachionus rubens and B. calyciflorus .
2. The effects of food quantity were well described by a modified Monod model with a threshold for zero population growth. The model parameters depended on particle size and nutritional quality of the food algae. Differences between the rotifer species were significant and reflected their varying food-size preferences.
3. For each rotifer species, thresholds were lowest for algae in the most readily ingested size range. The lowest thresholds were 0.07–0.09 mgC 1⁻¹ with algae of about 5 μm equivalent spherical diameter (ESD) for B. rubens , and 0.19 mgC 1⁻¹ with algae of about 10 μm ESD for B. calyciflorus .
4. Maximal growth rates ( r _max) were slightly below 0.8 day⁻¹ for both rotifers with most algal species. The highest r _max values for both rotifers were observed when Cyclotella meneghiniana was provided as food. With this alga, B. calyciflorus had a significantly higher r_max (1.02 day⁻¹) than B. rubens (0.838 day⁻¹).
5. From a comparison of the relationship between growth rates and ingestion rates, Chlamydomonas reinhardii appeared to be of low nutritional quality for B. rubens .
6. Egg ratios were related to growth rate and were not influenced by the algal food used. Egg development times and average mortality rates were estimated from the relationship between egg ratio and growth rate. B. calyciflorus appeared to have a high average mortality rate (0.383 day⁻¹) compared to B. rubens (0.083 day⁻¹).     

Life history and production of Paranephrops zealandicus in a forest stream, with comments about the sustainable harvest of a freshwater crayfish

1. Life history and production were assessed for the crayfish Paranephrops zealandicus in three reaches of a headwater stream with a catchment of regenerating coniferous-broadleaf forest in the south-east of the South Island of New Zealand.
2. Crayfish density ranged from 3 to 4 m⁻² in riffles and 4–12 m⁻² in pools, depending on reach. Crayfish biomass (4–33 g AFDW m⁻²) and annual production (2–11 g AFDW m⁻²) were high compared with values reported elsewhere, while P:B ratio was low (0.33–0.43). This substantial production was dependent primarily upon high biomass rather than high growth rate.
3. The crayfish of this population rank amongst the longest lived and slowest growing ever recorded. Individuals estimated to be 16+ year of age were not uncommon. Females became reproductively active at 6+ year. Fewer than 4% of females carried eggs, and young remained attached to females for at least 15 months.
4. We propose that characteristics of this population are the consequence of a cool thermal regime (mean daily stream temperature = 7.0 °C, range 1.8–11.9 °C), and that low biomass turnover and poor reproductive rate precludes any sustainable commercial harvest of crayfish from streams in New Zealand with similar thermal regimes.     

Influence of UV-B radiation on developmental changes, ethylene, CO2 flux and polyamines in cv. Doyenne d'Hiver pear shoots grown in vitro

In vitro shoots of cv. Doyenne ďHiver pear ( Pyrus communis L.) were irradiated under controlled environments for 6 h per day at 5 different levels of biologically effective UV-B radiation (UV-B_BE). UV-B exposure caused a progressive increase in apical necrosis above background levels and stimulated leaf abscission. Shoots grown for 2 weeks at 7. 8 mol m⁻² day ⁻¹ of photosynthetic photon flux (PPF) and treated with 8. 4 or 12. 0 kJ m⁻² day ⁻¹ UV-B_BE produced up to 4 times more ethylene than those given 2. 2 or 5. 1 kJ m⁻² day⁻¹ UV-B_BE or untreated controls. Exposure of shoots to 12 kJ m⁻² day ⁻¹ of UV-B_BE caused an increase in free putreseine content after 4 to 14 days of irradiation. Shoots showed a decrease in CO₂ uptake after 3 days of UV-B: thereafter, they appeared to recover their photosynthetic capacity. Under typical PPF conditions used in micropropagation (90 μmol m⁻² S⁻¹). 8. 4 kJ m⁻² day ⁻¹ of UV-B radiation was injurious to realatively tender tissues of in vitro pear shoots: increasing the level of UV-B_BE to 12 kJ m⁻² day⁻¹ produced even more adverse effects.     

Growth and growth substrate levels in spinach under non-steady state conditions of nitrogen nutrition and light

Spinach plants ( Spinacia oleracea L. cv. Subito) were grown in a complete nutrient solution under ample light intensity (14 h day⁻¹ at 660 μmol m⁻² s⁻¹) before being transferred either to a minus-N solution (experiment 1), or to limiting light conditions (6 h day⁻¹ at 220 μmol m⁻² s⁻¹; experiment 2). Shoot growth in experiment 1 decreased significantly from 0.24 day⁻¹ to 0.07 day⁻¹ after the fourth day of transfer. Root relative growth rate increased after 1 day from 0.25 to 0.31 day⁻¹, but decreased on the fifth day after transfer to 0.11 day⁻¹. Shoot growth in experiment 2 decreased significantly from 0.25 to 0.17 day⁻¹ after the fourth day of transfer, while root growth decreased to half of its original level (0.25 day⁻¹) already on the second day. Growth substrate levels in the plants (free sugars, free amino acids) and starch levels depended on the plant age, the moment in the diurnal cycle, and the imposed treatment. Fluctuations in shoot growth or root growth resulting from the light or N limitation could not be explained by a correspondent increase or decrease in the levels of growth substrates. The hypotheses underlying the functional equilibrium theory, assuming shoot and root growth to be controlled by N- and C-containing substrates respectively, and several other growth and partitioning models are therefore questioned. A neglect of the osmotic role of the free sugars in these models might be the explanation for this.     

Contribution of grazing in phytoplankton overall losses in a shallow French lake

SUMMARY 1. Based on data for ¹⁴C-primary production and biomass changes in a small and shallow lake (Créteil Lake, France), overall phytoplankton losses were calculated through an annual cycle (November 1985-October 1986). The summer period in 1986 is compared with two other summer periods in 1985 and 1980, these two years corresponding to extreme levels of algal biomass.
2. Independent from the trophic state of the lake, phytoplankton populations were dominated by small-sized species (<20 μm); their high growth rate (maximal in May and June: 0–8 day⁻¹) was characteristic of nanoplanktonic natural populations.
3. The positive correlation between phytoplankton losses and production indicates a close coupling between growth and loss processes.
4. With a high filtering rate (0.22 day⁻¹ as an annual average), zoo-plankton impact is considerable at any time of the year but especially in late summer, when grazing losses exceeded primary production.
5. Despite the uncertainty concerning the meaning of ¹⁴C-primary production, the persistence of small algae throughout the year implies that a great part of the phytoplankton production was harvested by grazers which led to a recycling of organic matter within the water column.