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Seasonal variations in the physiological condition of the benthic amphipods Monoporeia affinis and Pontoporeia femorata in the Gulf of Riga (Baltic Sea)
Kari K. Lehtonen
Finnish Institute of Marine Research (FIMR) P.O. Box 33, FIN-00931, Helsinki, Finland; (phone: +358-9-613 941; fax: +358-9-613 944 94; e-mail: lehtonen@ﬁmr.ﬁ)
Received 5 August 2002; accepted in revised form 4 February 2004
Key words: Ammonia excretion, Baltic Sea, Benthic amphipods, Gulf of Riga, Lipids, Seasonal variation
Abstract Seasonal variations in the physiological condition of two benthic deposit-feeding amphipods were studied in two environmentally differing areas in the Gulf of Riga ͑Baltic Sea͒ with the purpose of estimating their role in benthic mineralisation. At the shallower ͑26 m͒ nearshore station, Monoporeia affınis exhibited low ammonia excretion rate ͑VNHϩ 4 ͒ throughout the year. After intensive accumulation of lipids in spring, the lipid reserves of the individuals became rapidly depleted during autumn. Gravid females in winter-early spring showed exceedingly low lipid and triacylglycerol ͑TAG͒ levels compared to studies carried out in open-sea areas. Pontoporeia femorata studied at the deeper ͑43 m͒ offshore station exhibited a higher VNHϩ 4 at all times with no signiﬁcant seasonal variation. Lipid and TAG accumulated continuously until November, reaching higher levels than recorded previously for this species. Experiments showed that the temperature coefficient Q10 determined for VNHϩ 4 is likely to depend on the recent feeding history of the amphipods. Mineralisation of nitrogen and carbon by the –2 yr–1. amphipod populations at the study sites was estimated to be 1.7-3.0 g C and 11.3-13.5 mmol NHϩ 4 m Conclusively, both biotic and abiotic factors affect the condition of the amphipods, while the species also have genuine differences in their metabolic characteristics. clusively a subsurface deposit feeder ͑Byrén et al. 2002͒. Physiological characteristics of these species have been shown to reﬂect benthic nutritional conditions ͑Lehtonen 1994, 1995, 1996, a,b͒; variability in lipid level and lipid class composition indicated different bioenergetic strategies in amphipod populations inhabiting hydrographically different environments, while an elevated ammonia excretion rate ͑VNHϩ 4 ͒ is usually caused by recent feeding on highquality food. The Gulf of Riga is a semi-enclosed bay ͑mean depth 26 m͒ of the Baltic Sea, with water exchange with the Baltic Proper through relatively narrow straits ͑Figure 1͒. The gulf is greatly inﬂuenced by riverine input from a large drainage area, with the
Introduction The amphipods Monoporeia affınis ͑Lindström͒͑formerly named Pontoporeia affınis; see Bousﬁeld 1989͒ and Pontoporeia femorata Krøyer are usually highly dominant members of the soft-bottom macrozoobenthic communities of the northern Baltic Sea ͑e.g., Segerstråle 1937; Elmgren 1978; Andersin et al. 1978, 1984; Lehtonen and Andersin 1998͒. As deposit-feeders they are exceedingly dependent on the nutritional characteristics of the upper sediment layer, which, in turn, is governed by the quantity and quality of sedimenting particulate matter. Furthermore, M. affınis has been shown to feed mostly on the sediment surface, while P. femorata is almost ex-
playing an important role in benthic mineralisation and physical reworking of the sediment surface layer. seasonal variation in the physiological condition of M. femorata from two different stations was investigated by measuring VNHϩ 4 . river Daugava as the major source of nutrients and particulate matter. Köuts and Håkansson 1995͒ and primary production ͑Andrušaitis et al. a nonindigenous spionid polychaete. Cederwall et al. invaded the soft bottoms in the 1990s with an exceptional efficiency ͑Cederwall and Jermakovs 1999. 1996͒ were utilised to describe the benthic habitats. Marenzelleria viridis. Uitto and Sarvala 1991. Still. Since environmental factors have a profound effect on animal populations. Lipids play an integral role in the bioenergetics of M. Large spatial and interannual variability in amphipod population densities are well-documented features. these investigations serve to estimate the role of the amphipod species in the benthic mineralisation processes in the Baltic Sea benthic environments. JML. dissolved oxygen͒ and comparing these with results from studies in open-sea areas where these factors remain relatively stable throughout the year ͑Lehtonen 1995.g. 1987͒. but also to competition in overgrown populations ͑e. Since M. Moreover. Main study stations: GR1 and GR5. affınis inhabiting a relatively shallow marginal embayment that exhibits extensive seasonal variation in hydrographic conditions ͑temperature. GF1 and GF2. An elevation in benthic biomass. the feeding habits of the two species. lipid level and lipid class composition. 1999͒. Benthos samples for the description of community structure and population dynamics of the amphipods were also taken. hence. and these phenomena have mainly been attributed to variability in food resources. Maps of the Baltic Sea ͑A͒ and the Gulf of Riga ͑B͒. affınis and P. In this study. Andersin et al.442 Figure 1. afﬁnis and P.. Lehtonen and Andersin 1998͒. 1993. the population dynamics of the species. supplementary study stations for VNHϩ 4 studies: SR5. more information on their metabolic features is needed for a better understanding of their consumption of deposited organic material and. A gradual eutrophication of the gulf during the past decades has been observed ͑Ojaveer 1995͒. mostly due to an increase in the abundance of molluscs. M. The amphipod population data were combined with the VNHϩ 4 measurements and previous studies on respiration rates ͑Lehtonen 1996b͒ and used in es- . 1992͒. has also been recorded ͑Lagzdins et al. one of the main aims of the present study was to examine ecophysiological characteristics of M. 1996a. affınis and P. In addition. Parallel sediment studies ͑Carman et al. femorata remain as important members of the area’s benthos as food for higher trophic levels. b͒. generally. 1978. femorata and determine their reproductive output and. characterised by relatively high levels of nutrients ͑Yurkovskis et al. femorata occupy and feed in different sediment surface layers. affınis and P.
In most cases. as modiﬁed by Bayne et al. After a 24 h acclimatisation period 1-3 individuals were incubated in 23. Finnish model 0.5 mm sieves and stored in 5% formalin solution buffered with hexamin. 4 g dry wt The effects of temperature on VNHϩ 4 were studied by additional experiments carried out in May 1995. and GR5 ͑43 m͒ in the southwest-central basin of the Gulf of Riga ͑Figure 1͒.5-mm length classes. August. In addition. unpublished data͒. They were regarded suitable because of clearly differing physical environments. using cooled ͑4-6 °C͒ seawater from the respective sites. ͑1985͒. Selection of the stations was based on a benthic survey covering over 20 sites in the Latvian part of the gulf. Temperature coefficient ͑Q10͒ for VNHϩ 4 was determined using the equation log Q10 = ͓log10 ͑k2 ⁄ k1͔͒ / ͑t2 Ϫ t1͒ where t1 and t2 are the experimental temperatures ͑4 and 12 °C͒ and k1 and k2 are the excretion rates at the respective temperatures. and dissolved oxygen concentration by Winkler titration. affınis at Station GR1 and P. The amphipods ͑M.4 mm͒ aquaria situated in the cold laboratory ͑aboard ‘Aranda’͒ or refrigerator ͑aboard Latvian vessels͒ adjusted to 4 °C as used in the VNHϩ 4 experiments. July. Total lipid and lipid class analysis Freeze-dried amphipods were analysed for total lipid and lipid class composition using the micromethod of Gardner et al. with some modiﬁcations ͑Lehtonen 1995͒. only individuals from the reproducing generation ͑both species are semelparous͒ were studied. close to the mouth of the river Daugava. passed through 1. with 3-10 replicates and the excreted NHϩ 4 was determined using the method of Solórzano ͑1969͒. Unpublished data from similar experiments on Q10 of VNHϩ 4 of the two amphipod species. September and November 1994 and January. 5 van Veen samples were taken on each sampling occasion.1 m2͒ and passed through a small 1-mm sieve. adequate amphipod densities and for being within reasonable logistic distance from the shore for a year-round sampling scheme. NHϩ 4 –1 –1 excretion is given as mol NHϩ d . April. At each station.04 m2. Immediately after sampling the individuals were transferred to aquaria kept at 12 °C. ͑1999͒. enumerated and weighed ͑wet wt͒. Sampling was carried out with r/v ‘Aranda’ of the Finnish Institute of Marine Research ͑FIMR͒ and the Latvian vessels ‘Geofyzikis’ and ‘Antonija’ in January. femorata at Station GR5͒ were rapidly transferred to sand-bottom ͑grain size Ͻ 0. NH+ 4 excretion measurements excretion measurements were carried out NHϩ 4 aboard the vessels according to Lehtonen ͑1994. ͑cf. The individuals used in the experiments were stored at –80 °C or in liquid nitrogen and freeze-dried. The body length of the individual amphipods was measured from the tip of the telson to the end of the rostrum to the nearest 0.443 timating the contribution of the amphipod populations to mineralisation of carbon and nitrogen at the study sites. 1995͒. 1976͒ the animals were identiﬁed to species level. followed by gravimetric determination of total lipid and lipid class analysis by TLC-FID using silica-layered Chromarods ͑S-III͒ and Iatroscan MK-5 analyser. After a minimum of 3 months in storage ͑Dybern et al.0 and 0. 3-6 samples were taken with van Veen grabs ͑biting areas: Latvian model 0. Estimations of respiration and NH+ 4 release of the amphipod populations Amphipods from each sampling date were grouped into 0. considered valid also for specimens from the Gulf of Riga: Material and methods Sampling Seasonal studies were performed at two stations: GR1 ͑depth 26 m͒. Near-bottom salinity and temperature were recorded each time with a CTD sonde. Cederwall and Jermakovs ͑1999͒ and Cederwall et al. Mean wet weights of the individuals representing each length class were obtained using the following length-weight power function equations ͑FIMR. March and May 1995.9 ml ﬂasks in the dark at 4 °C for 10-14 h.5 mm. . and the VNHϩ 4 measurements were started after a 24 h acclimation period. Lipids are extracted from each individual in chloroform-methanol ͑2:1͒. ͑1985͒. performed at 4 deep ͑ Ͼ 80 m͒ open-sea stations in the Gulf of Finland and Bothnian Sea are also included here.
femorata has been shown to have a lower metabolic rate than M. b ϭ non-empirical exponent 0. af finis: W ͑mg wet wt͒ = 0. Application of M.770 ͑October-April period͒.29W ͑mg dry wt͒0. affınis data to P.75 was chosen to convert VO2 to carbon ͑C͒ units. VNHϩ 4 and VO2 were temperature-corrected using respective Q10 values of 3. The VNHϩ 4 of 1-mm length classes was calculated by ﬁtting the mean individual dry weights from each length class into the equation E ϭ aW0. Winter values for oxygen are not available.7 for M. showing the ϩ level of NHϩ 4 excretion ͑i. affınis ͑Lehtonen.7 and 2. affınis and P.75. determined for M. femorata: W ͑mg wet wt͒ = 0. M . Empirical Q10 values of 1. near-bottom temperatures up to 12 °C ͑GR1͒ and 9 °C ͑GR5͒ were recorded.018L ͑mm͒3. respectively. while during winter-spring the temperature remained between 0. Results Temperature and oxygen conditions In the summer. and 2.06 An empirical factor of 0. Seasonal variation in temperature ͑A͒ and dissolved oxygen ͑B͒ in the near-bottom water at the stations GR1 ͑solid line͒ and GR5 ͑broken line͒ during the study period. determined for M. and VO2 ͑l O2 d–1͒ ϭ 13. 1983͒ and a ϭ mean intercept value. Both excretion and respiration values were multiplied by the number of individuals in each respective length class. and the metabolism of the whole population was calculated by summing-up the values calculated for each length class.25 ͑Lehtonen and Andersin.05 P . Annual total mineralisation ͑as g C or 4 –2 mmol NHϩ m yr–1͒ was calculated using linear in4 terpolation between the values measured at each sampling date. 1996b͒ were used for both species. calculated for each sampling date.3- . the magnitude of mineralisation of C by this species in the study area is considered accurate using these equations. For the estimation of C mineralisation. W ϭ mean dry weight ͑mg͒ of the individuals from a given length class.e.0 for VO2 ͑Lehtonen. femorata. 1998͒ was applied to convert the wet weights to lyophilised dry weights.͒ were used for M.444 Figure 2. femorata.016L ͑mm͒3.8 and 1. However. femorata probably overestimates the mineralisation potential for the letter species since P. Detailed seasonal studies of this kind have not been performed on P. affınis. a ϭ VNH4 ͒. femorata. affınis ͑Lehtonen.. 1996b͒.32W ͑mg dry wt͒0.774 ͑MaySeptember period͒.0 for P. VO2 ͑l O2 d–1͒ ϭ 18.75 ͑Peters. Mean dry weights from each length class obtained this way were ﬁtted into the model E ϭ aWb –1 where E ϭ excretion rate ͑mol NHϩ 4 d ͒. the equations given above for VO2.030 ͑gravid females͒. affınis ͑Cederwall 1979͒. Since the amphipods use mostly lipids for metabolism a respiratory quotient ͑RQ͒ of 0. O2 consumption ͑VO2͒ of both amphipod species was estimated using equations VO2 ͑l O2 d–1͒ ϭ 14. where E ϭ –1 excretion rate ͑mol NHϩ 4 d ͒. Daily respiration and NHϩ 4 release are given as mg C or mol –2 –1 NHϩ m d . W ϭ lyophilised dry wt ͑mg͒ and a ϭ ‘date-speciﬁc’ intercept ͑Table 3͒.93W ͑mg dry wt͒1. in prep.8 and 2.
2 87.1%͒ ͑Figure 4͒. M. Oxygen concentrations showed signiﬁcant seasonal variation ͑3 to 12 mg O2 l–1͒ with a longer period of low-oxygen conditions at Station GR5 in late autumn ͑Figure 2͒. a 3-value moving average was used and the individuals were divided into year classes ͑cohorts͒.8-7. while the bottom at Station GR5 was muddy and had a high organic content ͑Table 1͒. affınis͒ and 3. except mean grain size ͑Pallo et al. At Station GR1 a high abundance and biomass of the deposit/ﬁlter-feeding clam Macoma balthica ͑1429 ind m–2. femorata͒ lyophilised dry wt.7 41. The community data ͑abundance and biomass͒ are presented as annual means ͑Figure 3͒. 1998͒. Corophium volutator was present ͑170 ind m–2͒. however. At Station GR1. ͑1996͒. 55% of total biomass͒ was observed. 12% of total biomass͒. The amphipod populations At Station GR1.7 13. affınis was not studied at Station GR5.7%͒ compared to 3 July ͑36.3 2. Sediment characteristics and benthic community structure The study sites differed markedly in terms of sediment characteristics and benthic community structure. For size distributions. with complete absence of P. At Station GR1 the annual average of total abundance of macrozoobenthic organisms was an order of magnitude higher compared to Station GR5 ͑5657 and 499 ind m–2.1 mg ͑P. the bottom consisted of muddy sand with a low organic content of the top sediment layer. femorata was 10 times higher in abundance compared to the co-existing M. affınis͒. respectively͒. also another benthic amphipod. The high density of the polychaete M. Station GR1 26 m Sediment description Mean grain size ͑m͒ – 0-1 cm Loss on ignition ͑%͒ – 0-1 cm – 1-2 cm Water content ͑%͒ – 0-1 cm – 1-2 cm Organic C ͑mol g dw–1͒ – 0-1 cm – 1-2 cm Organic N ͑mol g dw–1͒ – 0-1 cm – 1-2 cm Sandy aleurite.4 mg ͑M.8 7. Here. 3. light brown down to 2.445 Table 1.13. light brown down to 4. Sediment characteristics at the sampling stations in June 1991. affınis. At Station GR5 P. The weight ranges ͑obtained from monthly means͒ of the amphipods studied were 2. affınis at Station GR1 showed a slight decline by 14 September ͑31. femorata͒ or March-April ͑M.7% of total biomass͒.7 71. respectively͒ while the biomass was four times higher ͑156 and 38 g wet wt m–2͒ ͑Figure 3͒.0 84.5 cm 6.8 Station GR5 43 m Sandy-mud. femorata. Due to the difficulty in obtaining sufficient numbers of specimens at every sampling occasion. viridis at Station GR1 ͑2374 ind m–2͒ represents 33% of the total biomass.0 °C at both stations ͑Figure 2͒.7 14. At Station GR5 this species plays a less signiﬁcant role ͑62 ind m–2. affınis was the dominant amphipod species ͑1454 ind m–2.0 cm 11. By 11 November. the . both showed low densities ͑374 and 34 ind m–2. All data from Carman et al.0 2576 878 3950 3422 200 61 380 348 1. Both species exhibited a 2-year lifecycle and offspring were released in late January ͑P. M. Lipid level and lipid content The lipid levels of M.
femorata ͑5. the ‘lipid balance’ of P. affınis between 3 July and 11 November ͑Table 3͒.. in addition to presenting the proportion of lipids of total dry weight ͑Figure 4͒.2%͒. exhibiting low lipid levels on 30 March ͑9. The most signiﬁcant differ- .3%.8%͒ ͑Figure 5͒.e. affınis. affınis at Station GR1.2%͒ ͑Figure 4͒. during maturation. sterols/diacylglycerols and acetone-mobile polar lipids͒ comprised altogether 6. net depletion of lipid intensiﬁed towards late autumn. affınis was found to be size-dependent ͑Table 2͒. Seasonal variability in the lipid content of a ‘standard-size’ P.7 and 21.0–80. The other measured lipid classes ͑wax esters. After pooling the seasonal data. Macrozoobenthic community structure at the study stations ͑GR1 and GR5͒.5–20. Contrary to M. affınis͒ was not marked until 10 November.0% in P.. affınis. the lipid level of the ‘new’ reproducing generation ͑born 1994͒ was high ͑37. Calculations based on changes in mean weights and lipid levels showed no net accumulation of lipid in M. while the lipid content of the gravid females ͑5 January͒ was signiﬁcantly lower compared to others ͑Table 2͒. The dependence of the TAG content ͑mg ind–1͒ on the dry weight of the individuals as well as seasonal variability in the content of TAG were very similar to those of total lipid in both species ͑Table 2͒. Size-corrected values of the lipid content of the ‘new’ reproducing generation ͑born 1994͒ on 6 May 1995 were not different from previous year’s summer values. free fatty acids.8 and 38. between 17. and. obtained in a similar way as described for M. the lipid content ͑mg ind–1͒ of M. seasonal variability in the lipid content was examined also by comparing the lipid contents of a ‘standardsize’ individual.8%͒. the proportion of phospholipids ͑PL͒ varied between 10.7 mg dry wt.1–15. NH+ 4 excretion: seasonal variation A power failure in the ultrafreezer ͑ Ϫ 80 °C͒ caused the destruction of the samples collected in January lipid levels had declined to 20. data from March and May 1995 not available͒.4% and 7.9%. lipid content regression equation given in Table 2 and the mean value of each monthly average weight of individuals representing the reproducing generation ͑2. using the dry wt vs. Gravid females on the brink of spawning on 5 January showed a lipid level of 20. Thus.3%͒ and was still relatively high on 11 November ͑30. Instead.446 ence was found between the gravid females and the specimens collected on all other dates.7 mg dry wt. femorata. net accumulation͒ throughout the autumn.6% and. presented as average abundance ͑above͒ and biomass ͑below͒.8%͒ compared to M. femorata was positive ͑i. and the gravid females showed a markedly high depletion rate by 5 January. In P. ovigerous/gravid females continued to deplete lipid ͑5 January: 16.1%͒. Lipid class composition In both species the seasonal patterns in triacylglycerol ͑TAG͒ levels followed closely those of total lipid levels. Figure 3.3–37. The lipid level of P. turning negative ͑i. affınis at Station GR1 ͑44.e. femorata at Station GR5 remained almost constant throughout the summerautumn ͑34. in M. P. affınis was signiﬁcantly reduced by 11 November. femorata and M. femorata at Station GR5 showed constantly higher TAG levels ͑seasonal range: 63. net depletion͒ only between gravidity and offspring release ͑Table 3͒.3%. After the spring bloom in early May 1995. The lipid content of M. respectively.5–72.
affınis ranged be–1 dry wt d–1 durtween 13. While acknowledging these results. enabling a larger weight scale than possible with ‘single-date’ measurements. affınis. VNHϩ 4 power regression proved to be non-signiﬁcant ͑F1. excretion rate data. 40 ϭ 4 d ͒ ϭ 0. r ϭ 0. femorata. For M.79. 32 ϭ 7. estimations of the mean dry weights of the specimens used in excretion experiments were obtained from population samples and used for the calculation of VNHϩ 4 .9. and.447 Figure 4. ANOVA performed on weight-corrected rates calculated for a ‘standard-size’ individual of 2. August ͑not at GR1͒. p ϭ 0. These factors include individual differences in recent feeding history or the amount of energy reserves which are likely to cause more inter-individual variability in VNHϩ 4 than in oxygen consumption.55 tion E ͑mol NHϩ ͑F1. p Ͻ 0.0͒ but is affected also by other factors ͑low r2͒.75 ͑Peters 1983͒ was considered in this case to be the most reliable method in examining the differences between the sampling dates and stations.0004͒.0. further. However.4. Seasonal variation in lipid levels ͑% dry wt͒ of Monoporeia affınis and Pontoporeia femorata. and this data was omitted as well as the signiﬁcantly higher rates recorded on 6 May during high sedimentation of the spring bloom at Station GR1. November 1994 and May 1995 for both stations separately.054W ͑mg͒ 2 17. July. 22 ϭ 0. The 4 males caught at Station GR5 on 11 November showed signiﬁcantly higher VNHϩ 4 compared to others ͑subadults͒͑two-sample t-test on weight-corrected rates: df ϭ 10. Table 4͒. the equa–1 0.39͒. At Station GR1 the VNHϩ 4 of M. giving evidence that the VNHϩ 4 is clearly weight-dependent ͑slope deviating signiﬁcantly from 1. p ϭ 0. For P. the data were compiled selectively. Actual weights were not available on 19 April. Because of an attempt to establish a weight-rate relationship not affected by a recent input of goodquality food ͑spring bloom sedimentation͒ or physiological modiﬁcations related to reproduction. t ϭ 2. with the highest mean rate recorded on 6 May 1995 ͑Figure 6. weight-correction of the excretion rates using the common scaling exponent 0. the dry weight vs. Tukey’s test showed that . Similar to lipid and TAG contents.05͒. the VNHϩ 4 is given both as direct values per weight and as per ‘standard’ individual weight for the whole study period. p Ͻ 0.9 mol NHϩ 4 g ing the season.31͒ was determined. Results from January and March 1995 experiments were omitted from this analysis because of the small number of measurements ͑due to difficulties in sampling͒. Vertical bars indicate SE unless smaller than symbol. Gravid females were also discarded because of a distinctly higher VNHϩ 4 which is unrelated to food conditions ͑Lehtonen 1996b͒.7 mg showed a significant difference in VNHϩ 4 between the sampling dates ͑F4.0001. and April 1994. The effect of body weight on VNHϩ 4 of each species was studied by pooling the seasonal weight vs.0 and 28. These were not included in the analyses because of a higher activity probably related to increased swimming activity and/or metabolic adjustments related to reproduction. ANOVA was performed on VNHϩ 4 measurements made in April. September. in the estimations of the mineralisation potential of the populations.
01*. femorata was recorded at Station GR5 on 6 May 1995. df ϭ 17. femorata ͑standardised for a 5.426 . at that time. 61 56 43 56 290. p ϭ 0.2 81. 42 ϭ 1.001.9͒ was calculated for individuals collected from Station JML ͑western Gulf of Finland͒ on 24 April 1996 ͑4 °C: 16.05͒.216 dry wt ͑mg͒ Ϫ 0. At Station GR5.455 .129 dry wt ͑mg͒ Ϫ 1. 11 November: p Ͻ 0. Seasonal changes were studied using values calculated for a ‘standard-size’ individual of each species.05 and ns ϭ not signiﬁcant. the highest coefficient ͑3.0001 0.11͒.046 . p ϭ 0. ANOVA Monoporeia affınis ͑GR1͒ Lipid ͑mg͒ ϭ 0.seasonal variation: ANOVA Tukey’s multiple comparison test Monoporeia affınis ͑2.448 Table 2. the range being 20. ANOVA without data from 6 May showed no difference between the rates ͑F3.0001 0.8 18.0001 0.7 mol g dry wt affınis. ** ϭ p Ͻ 0.332 dry wt ͑mg͒ ϩ 0. 36 24 30 23 F1. * ϭ p Ͻ 0. In the Tukey’s multiple comparison test Table created for lipid and TAG ͑in parentheses͒.0 Ͻ Ͻ Ͻ Ͻ Ͻ Ͻ Ͻ Ͻ 0.36. Table 4͒.19͒. The lowest value for Q10 ͑1.7-30.0001 17 Aug 14 Sep 11 Nov 05 Jana 30 Mara – – – – *** ͑*͒ *** ͑**͒ *** ͑***͒ ns ͑ns͒ * ͑ns͒ *** ͑**͒ *** ͑**͒ ns ͑ns͒ *** ͑**͒ *** ͑***͒ Pontoporeia femorata ͑5.4.1 15.6 mol g dry wt–1 d–1͒.1 62.0001 0. seasonal variation in the VNHϩ 4 of P. a ϭ gravid females.8͒ for P. Monoporeia affınis and Pontoporeia femorata. when the –1 –1 d .24͒. .7 mg dry wt͒ Date 03 Jul 03 Jul 14 Sep 11 Nov 05 Jana 30 Mara 06 May ns ͑ns͒ *** ͑***͒ *** ͑***͒ *** ͑***͒ ns ͑ns͒ F df p ϭ ϭ ϭ ϭ ϭ ϭ ϭ ϭ 24.2͒ was recorded on 20 September 1995 at Station GF2 ͑central Gulf of Finland͒ for P.seasonal variation: ANOVA Pontoporeia femorata ͑GR5͒ Lipid ͑mg͒ ϭ 0.7 mg individual͒ was not signiﬁcant ͑ANOVA: F5. except on 14 September ͑p Ͼ 0.05͒. The highest Q10 ͑2.001.7 mg dry wt͒ Date 02 Jul 17 Aug 02 Jul 17 Aug 14 Sep 10 Nov 05 Jana 30 Mar 06 May ns ͑ns͒ ns ͑ns͒ *** ͑**͒ *** ͑***͒ ** ͑**͒ ns ͑ns͒ 14 Sep 11 Nov 05 Jana 30 Mar ns ͑ns͒ * ͑ns͒ *** ͑***͒ ns ͑ns͒ ns ͑ns͒ * ͑ns͒ *** ͑***͒ ns ͑ns͒ ns ͑ns͒ *** ͑***͒ ns ͑ns͒ * ͑ns͒ *** ͑**͒ *** ͑***͒ ns ͑ns͒ the rates measured on 6 May 1995 were higher than on all other dates ͑20 April and 2 July: p Ͻ 0.0001 F1. For M.0001 0.0 0.7 mol g dry wt–1 d–1͒ and lowest ͑2. at a time when no signiﬁcant difference could be noted in VNHϩ 4 measured either at 4 or 12 °C ͑t-test: t ϭ 1. F5. VNHϩ 4 at 4 °C was 32. F6. 26 ϭ 2.seasonal variation: ANOVA TAG ͑mg͒ ϭ 0. VNHϩ 4 at 4 °C was as high as 64.2͒ at the same station on 13 August 1996 ͑4 °C: 26.0 6. F1.282 dry wt ͑mg͒ ϩ 0. femorata.6 mol g dry wt–1 d–1. F6.02 0. Statistical analyses on the lipid and triacylglycerol ͑TAG͒ content of the individuals.3 11.211 . statistical signiﬁcancies are indicated as *** ϭ p Ͻ 0.24.seasonal variation: ANOVA TAG ͑mg͒ ϭ 0. NH+ 4 excretion: effect of temperature on rates The results indicate that the magnitude of the effect of temperature on VNHϩ 4 is dependent on the initial level of excretion ͑at 4 °C͒ ͑Figure 7͒. –1 p ϭ 0. F5. F1.3 mol NHϩ 4 g –1 dry wt d ͑Figure 6.
TAG ϭ triacylglycerols. Seasonal variation in lipid class composition of Monoporeia affınis and Pontoporeia femorata as % of total lipids ͑A͒ and weight ͑mg͒ of each lipid class in a mean-size individual ͑B͒. Other ϭ wax esters ϩ free fatty acids ϩ sterols/diacylglycerols ϩ acetone-mobile polar lipids. Vertical bars indicate SE unless smaller than symbol. data from March and May 1995 are missing because population samples needed for the calculation of mean weights were not collected. Results for April have been calculated using the mean weights of the studied generation obtained from population samples ͑see text͒. .449 Figure 5. Variation in the VNHϩ d ͒ for the reproducing generation of Monoporeia affınis and Pontoporeia femorata during 4 ͑mol NH4 g the study period. G ϭ gravid females. In ͑B͒. Note different scales in ͑B͒. PL ϭ phospholipids. ϩ –1 –1 Figure 6.
450 Table 3. and annual NHϩ NHϩ 4 m 4 –2 release was 11.0297 ͑0.0320 ͑0.0392 ͑0. Date 1994 Jan 27 April 19 July 02/03 Aug 17 Sep 14 Nov 10/11 . determined for the equation E ϭ aWb ͓E –1 ϩ ϭ mol NHϩ 4 d . and annual C mineralisation was estimated at 3.9 ͑May͒ to 67.0903 ͑0. The intercept values were used in the calculations of total NH4 release by the amphipod populations. affınis. Monoporeia affınis and Pontoporeia femorata.8 Ϫ 2.0470 ͑0.0 4. Offspring release occurred in March.7 ͑May͒ and 9. and by calculating a mean daily rate for each period.0023͒ 0. In M.75͔.Gravid females May 06 a n – 6 9 – 7 9 – 3 3 8 b GR1 Monoporeia affınis – 0. annual total being 1. resembling that recorded in the Bothnian Sea ͑Lehtonen 1996a͒ where a highly concentrated seasonal sedimentation pattern prevails.8 ͑July͒ mol NHϩ 4 m –2 ϩ annual NH4 release was 13. The seasonal range in lipid levels was wide ͑22-37%. 1996a͒ but the decline towards winter was more striking. 1992.0 g C m–2 yr–1.9 ͑January͒ and 15.0119͒ 0. 4 m Daily respiration ranged between 3.1 Ϫ 0. Lehtonen 1995.2%͒. the dependence of lipid levels on resource .1 Lipid level of M.0 ͑July͒ mg C m–2 d–1.0208 0.0487 ͑0.0189͒c 0.0223 – 0.1 Ϫ 14.4 Ϫ 0. affınis ͑GR1͒ 03 Jul Ϫ 16 Auga 16 Auga Ϫ 14 Sep 14 Sep Ϫ 11 Nov 11 Nov Ϫ 05 Jan ͑G͒ a Lipid g d–1 Ϫ 0.5 mmol NHϩ yr–1. femorata ͑GR5͒ 02 Jul Ϫ 17 Aug 17 Aug Ϫ 14 Sep 14 Sep Ϫ 10 Nov 10 Nov Ϫ 05 Jan ͑G͒ Lipid g d–1 6. affınis at Station GR1 were intermediate compared to other areas of the northern Baltic Sea ͑Hill et al.0120͒ 0.0270͒c 0.0054͒ 0. Net lipid accumulation or depletion rates ͑g d–1͒ in the reproducing generation.0081͒ ͑0. a ϭ intercept. similar to most areas of the northern Baltic Sea ͑Leppänen 1988.9 ͑April͒ –2 –1 d ͑Figure 8͒.0452 ͑0. Seasonal values of the intercept a ͑ Ϯ SD͒. affınis in 16 August used for calculation was estimated assuming a linear change between 3 July and 14 September Table 4. Discussion Seasonal variations in lipid levels and lipid class composition In summer.0118͒ 0. G ϭ gravid females.3 7.0078͒ 0.0056͒b ͑0. Heiskanen and Kononen. obtained by linear interpolation of changes in dry weight and lipid level ͑% dry wt͒ between sampling dates.0289 – n 7 6 9 8 6 3 3 4 3 10 GR5 Pontoporeia femorata 0.0164͒ ͑0.4 mol –2 –1 d ͑August͒͑Figure 8͒. femorata population at Station GR5 ranged from 3.0317 0. C mineralisa4 m tion was calculated to range between 0.0080͒ 0. 1994. describing the weight-speciﬁc NH4 exϩ cretion rate of the amphipods.5 P. Used for calculations in August 1994. and the gravid females were notably lipid-poor at that time ͑9.0105͒ Used for calculations in May 1994.0611 ͑0.0043͒ 0. c Used for calculations in December 1993 Mineralisation of N and C by the amphipod populations At Station GR1.0459 ͑0.0172͒a 0. b ϭ ﬁxed non-empirical exponent 0.3 mmol NHϩ yr–1. The release of NHϩ 4 by the P. lipid and TAG levels of M.0450 ͑0.9 mg C m–2 d–1 ͑August͒.0511 ͑0.0146͒ 0.7 g C m–2 yr–1.0792 ͑0. Monoporeia affınis and Pontoporeia femorata. M. gravid females not included͒. affınis population was estimated to range from 10.Males 1995 Jan 05 . while to 84. Lehtonen and Andersin 1998͒. n ϭ number of determinations. W ϭ dry weight ͑mg͒. the release of NHϩ 4 by the M.0051͒a ͑0.0390 ͑0.
A͒ and carbon mineralization ͑mg C m–2 d–1. Stations: GR1 and GR5 – Gulf of Riga. NHϩ d . GF1 and GF2 – Gulf of Finland. –2 –1 ϩ Figure 8.451 Figure 7. SR5 – Bothnian Sea. B͒ by Monoporeia affınis and Pontoporeia femo4 release ͑mol NH4 m rata populations at stations GR1 and GR5. . The rates have been calculated separately for each 1-mm size class but are shown here as combined to 2-mm size classes. as determined by VNHϩ 4 at 4 and 12 °C. Relationship between VNHϩ 4 ͑at 4 °C͒ and temperature coefficient Q10 for Monoporeia affınis and Pontoporeia femorata. JML.
femorata seems typical for this life stage. affınis at Station GR1 in late autumn may reduce the ability of the gravid females to survive and take care of their brood until hatching. Lehtonen 1996a͒. An even more marked elevation in the VNHϩ 4 after the spring bloom was noted in a deep.452 input and competition during the previous season was examined by Lehtonen and Andersin ͑1998͒. femorata at Station GR5 was higher than recorded by Paradis and Ackman ͑1976͒. affınis a complete halt in growth in early autumn has been recorded ͑Lehtonen and Andersin 1998͒. and/or the increasing excretion of the developing embryos within the female’s marsupium. affınis ceases much earlier ͑Uitto and Sarvala 1991͒. high VNHϩ 4 is not associated with food availability but caused by an increased use of body protein for metabolic energy production due to already depleted lipid reserves. femorata continuing to grow in autumn while the growth of M. Table 2͒ recorded for P. Male P. For comparison. the seasonal range in lipid levels was narrower than in M. The higher and seasonally more stable excretion rate of P. femorata showed signiﬁcantly higher VNHϩ 4 compared to subadults. and has also been observed in gravid M. and unpublished data͒ and is probably related to different feeding patterns and growth dynamics ͑Uitto and Sarvala 1991͒. previous studies on the ecological physiology of the species in other sea areas may be used to cast light on the underlying causes of the observed patterns in the condition of both amphipod species. In this case.0 g lipid d–1. Also in an open-sea population of M. Olli and Heiskanen 1999.3%͒ in P. affınis. In addition to functioning as stored energy for future metabolic needs lipids act as fuel for biosynthesis in the build-up of reproductive tissue and the formation of reproductive products ͑e. Reigstad et al. Potential effects of environmental conditions and intra/interspeciﬁc competition Because the two species were not studied in one and the same area. However.1 and 7. affınis at Station GR1. In 1995. the present study shows that P. with P. 1985͒. which is reﬂected in accelerated VO2 ͑Lehtonen 1996a. Falk-Petersen et al. Hill et al. Due to the hydrodynamic conditions prevail- . affınis at Station GR1 are characteristically found in poor-nutrition situations ͑e. Lehtonen 1995͒. femorata at Station GR5 between 2 July and 11 November differs markedly from that of M. affınis ͑Lehtonen 1996b͒. unpublished data͒. afﬁnis ͑Lehtonen. in a coastal area the two species have been shown to have different growth patterns.g. 1999͒. b͒.. corroborated by recordings of a lower swimming activity and VO2 ͑Cederwall 1979͒ in this species. 1992. a direct assessment of the effects of environmental setting on the physiological condition of the amphipods cannot be made. affınis population from the Bothnian Sea ͑Lehtonen 1996b͒. seek actively for females ͑Segerstråle 1937͒. after maturation. which has also been observed in male M. ͑1992͒ and Lehtonen ͑1995͒. which had a zero-or-negative lipid balance throughout this period. The reduced levels of lipid and TAG in gravid females seem characteristic for both species ͑Hill et al. The different lipid dynamics observed are likely related to differences in growth. while the signiﬁcantly higher value observed in May 1995 is considered a metabolic response to the sedimentation of fresh organic matter ͑Lehtonen 1996b͒. 1981. Seasonal and spatial variations in VNH+ 4 The low VNHϩ 4 levels recorded for M. in the open-sea areas of the northern Baltic Sea between late autumn and early spring͒. affınis mostly showed a negative lipid balance in late autumn ͑Lehtonen 1996a͒. Clarke et al. open-sea M. The highest mean lipid level ͑2 July 1994: 37. A higher VNHϩ 4 recorded here for gravid P. an open-sea population of M. Excluding gravid females. The greatly diminished lipid stores found in M. This may be caused by a general increase in metabolic activity which is connected to enhanced swimming as the males.. affınis ͑27-37%͒. A low lipid content of the offspring may also directly reduce their viability ͑Ouellet et al. affınis ͑see Hill et al. femorata have been interpreted as an implication of a different bioenergetic strategy compared to M. However. the peak of diatom sedimentation in the Gulf of Riga started to decline in the beginning of May. 1999.g. and are associated with increased protein metabolism. femorata seems typical for this species ͑Lehtonen 1995. The positive lipid balance ͑between 4. 1992͒. 1992. femorata is capable of attaining similar lipid and TAG levels often recorded for M. The lower lipid levels observed in P. and from thereon the settling material had greater proportions of heterotrophic biomass and slowly-sinking phytodetrital material ͑Lundsgaard et al.
The period of low oxygen concentrations observed in late autumn ͑Figure 2͒ may have affected the distribution and abundance of the amphipod species especially at Station GR5. It is therefore concluded that. 1998͒.and interspeciﬁc competition for food. In M. This important compartment of the benthos also utilises a signiﬁcant share of the annual input of quality food. the abundance of harpacticoid copepods of 140000 ind m–2 recorded at Station GR1 is among the highest observed anywhere in the world.g. Thus. the initial levels of VNHϩ 4 . and the overall higher diversity and biomass ͑Figure 3͒ at Station GR1 keeps the competition pressure at a markedly higher level as compared to Station GR5. reproduction and recruitment of amphiphod species. this material is of considerably lower nutritional quality as compared to a marine food source such as sedimenting algae.. balthica and C.. affınis ͑Ólafsson and Elmgren 1991. femorata ͑Figure 7͒. . This may be caused either by interspeciﬁc differences or. The inﬂuence of the river Daugava. 1982͒. as suggested by the apparent relation between the ‘initial’ excretion rate at 4 °C and Q10. while M. vascular plant and macroalgae remains͒ is high. The benthic community at the study sites differs in terms of species biomass and abundance and is likely to be affected by intra. especially M. Station GR5͒. showing that these amphipods are capable of reducing VNHϩ 4 in situations of a signiﬁcant loss of nitrogen. However. there exists evidence that meiofauna and even juvenile M. 1999. which is readily assimilated by the benthos ͑e. femorata at Station GR5 was not particularly high. Effects of temperature on VNH+ 4 The effect of temperature on the VNHϩ 4 was generally more pronounced for M. balthica are consumed by M. it is likely that the effect of temperature on VO2 is rather conservative ͑with Q10 values close to 2. while VNHϩ 4 is considerably more affected by nutrition. Ejdung et al. volutator. Importantly. the Q10 for VNHϩ 4 ranged between 2.g. and especially after the breakdown of the thermocline in autumn. Besides competition. The meiofaunal communities at both stations belong to the most abundant recorded in the Baltic Sea. Reigstad et al. in that study. indicating that temperature elevation increases the immediate use of protein for metabolic energy if Q10 for VO2 would remain around 2. a common mechanism may be at work in which pelagic eutrophication provides the benthos with increasing amounts of food. Parsons et al. 7-fold compared to the pre-bloom situation. Despite their apparent good physiological condition the abundance of P. affınis than for P. Similar phenomena have been recorded for zooplankters ͑reviewed by Le Borgne 1986͒. 1996͒. a constant provider of organic particulate matter of terrestrial origin ͑e.9.. Compared to Station GR5. affınis.0. 1977. affınis ͑Lehtonen 1996b͒. showing different Q10 values for v and VNHϩ 4 . by differences in environmental nutritional conditions. The high abundance of other macrobenthic deposit. femtemperature-independent VNHϩ 4 orata at Station GF2 during a high-excretion period ͑Figure 7͒ would in this case result in O:N ratios indicative for a shift towards a more lipid-based energy metabolism following temperature increase.and/or ﬁlter-feeders.2 and 3. affınis at this location occurred in very low numbers ͑Figure 3͒. in the deep areas of the Gulf of Riga.0͒. Since VO2 is mainly a measure of general metabolic activity.453 ing in the Gulf of Riga the sedimented matter is also effectively resuspended ͑Floderus et al. No deﬁnite conclusion on the effect of competition can be drawn here due to the lack of studies in similar habitats with differing community structures and densities nor a proper assessment made of the quantity and quality of particulate matter received by the different areas in the Gulf of Riga. while their number at Station GR5 is considerably smaller ͑35000 ind m–2. Tenore et al. Carman et al. but. This substantial transport of material from the basin margins may offer a more long-lasting food source for benthic organisms at Station GR5 as compared to Station GR1. 2000͒. At the near-shore Station GR1 the organic content of the sediment surface layer ͑0-2 cm in depth͒ was low ͑Table 1. 1998͒. 1999͒. the 10-fold higher macrofaunal density at Station GR1 ͑Figure 3͒ competing for the food resources is likely to have an effect on the physiological condition of species in this area. the acceleration of VNHϩ 4 was ca. Pallo et al. seem to determine the type of response caused by elevations in temperature. indicating also a balance of metabolic substrates at the ambient temperature. with densities of Ͼ 900000 ind m–2 ͑Pallo et al. A slight increase in VO2 ͑by 22%͒ after the sedimentation of the spring bloom has been recorded for M. the resuspended ﬁne particulate material is to a great extent transported towards the central-basin accumulation area ͑e.g. The almost recorded for P. The subsequent deterioration of oxygen conditions lead to a reduced survival.
mineralisation is very low during the winter-spring period ͑Figure 8͒.0 g C m–2 yr–1͒ for P.3 °C. femorata may have different capabilities of modifying their bioenergetic strategy and acclimatisation to varying temperature and feeding conditions. however.454 The ‘temperature history’ of populations may play a role in adaptation. affınis. a signiﬁcant elevation in the metabolism of the population is observed.2 °C. femorata and M. The estimated daily NHϩ 4 release values ͑3. affınis and P. JML 1996: 3. GF1 and GF2͒ the populations in the Gulf of Riga are subject to a signiﬁcantly greater seasonal amplitude in temperature ͑Figure 2͒. had a wider range with periods of 18 °C and the density of the population was considerably higher compared to the present study.4-5. Lehtonen and Andersin ͑1998͒ showed the annual respiration of the M. ͑1999͒. further investigations on their ecophysiology and behaviour would be advantageous in understanding benthic nutrient dynamics in soft-bottom areas of the Baltic Sea.7 and 3. seasonal range 1991: 2. affınis population ͑11. Lehtonen. Between September 1991 and March 1992 in the Gulf of Riga. ͑1987͒ estimated the excretion of an abundant freshwater amphipod ͑Diporeia hoyi͒ population in Lake Michigan to comprise up to 42% of total benthic nitrogen release. Calculating the potential share of amphipod excretion of these rates gives values ranging between 3-13% of total NHϩ 4 efflux. Unfortunately. Considering that M. the late spring-early autumn period that presumably has the highest benthic activity was not studied by Aigars and Andrušaitis ͑1993͒. FIMR. 15-31 mmol –2 yr–1͒ in an area where the abundance and NHϩ 4 m biomass of the M. Annual rates ͑11. found M. the growth of the amphipods is intensive and together with a marked elevation in temperature. Compared to the amphipods at the deep stations in the Gulf of Bothnia ͑SR5͒ and Gulf of Finland ͑JML. The estimated annual total respiration ͑1. affınis population to be 14-17 g C m–2 yr–1. Aigars and Andrušaitis ͑1993͒ recorded total benthic ammonium efflux rates ranging between 30–2 –1 790 mol NHϩ d ͑mean ca.3 °C͒. and –2 –1 between 30-660 mol NHϩ d ͑mean ca. affınis to contribute 5-10% to the total NHϩ 4 efflux. .9 g C m–2 yr–1. 1994͒. Acknowledgements This study was part of the interdisciplinary multinational ‘Gulf of Riga Project’. femorata inﬂuence benthic mineralisation processes directly by metabolising organic material and also indirectly by bioturbation. in a laboratory experiment using manipulated sediment cores.7-4. nutritional factors seem to overshadow the effects of temperature on the VNHϩ 4 of these amphipods. with the exception of occasionally high values of ca.45. who. assuming benthic efflux rates at least twice higher than the higher mean would yield values between 4-10% for the contribution of the amphipods to total NHϩ 4 production.4 °C. ͑1983͒ showed that excretion by the amphipod C. Therefore. Bioturbation by C.5 mmol NHϩ 4 m corded by Lehtonen and Andersin ͑1998. open-sea station with a constantly low seasonal temperature range ͑2.9 to –2 –1 are average compared to m d 84. affınis and P.8 mol NHϩ ͒ 4 other sites studied in the Baltic Sea ͑up to 237 mol –2 –1 d . M. is much less than recorded by Bergström and Sarvala ͑1986͒ for a coastal M. 400 4 m –2 –1 ϩ mol NH4 m d ͒ at a 37-m accumulation bottom well comparable to Station GR5. affınis population is considerably higher but the VNHϩ 4 of the amphipods remains very low between autumn-spring ͑Lehtonen 1996b͒.3NHϩ 4 m –2 yr–1͒ are smaller than those re13. 31% of total benthic respiration͒. At the deep stations the seasonal temperature conditions were almost identical ͑Station SR5. GF2 1996: 3. Pelegrí et al. but is also inﬂuenced by temperature. In summer-autumn. Henriksen et al. Gardner et al. Apparently. Mineralisation of C and N by the amphipod populations The C and N mineralisation potential of the amphipod populations studied is mostly dependent on their abundance and biomass. It is therefore concluded that excretion by the amphipods plays a small but stable role in the benthic nitrogen dynamics of the Gulf of Riga. Temperature regime in that area.7-4. 30% at the times of very low total efflux rates –2 –1 d ͒ as recorded in January-Feb͑30 mol NHϩ 4 m ruary. funded mainly by the Nordic Council of Ministers. Partly due to the low temperature. These values are in accordance with the results of Tuominen et al. respectively. 300 mol NHϩ 4 m 4 –2 –1 m d ͒ at a 27-m station close to Station GR1. volutator has been shown to enhance also benthic denitriﬁcation rates ͑Pelegrí and Blackburn 1994. providing nitrifying bacteria a potentially signiﬁcant source of NHϩ 4 . 1995͒. unpublished data͒. At a deep. volutator could account for 80% of the net NHϩ 4 ﬂux from the sediment.
. ͑eds͒. pp. and Pallo P. Sars͒.. Burns K. Publication No. Arch.R. ICES J. and Malczyk J. Aigars J. Cederwall H. Baltic. Other members of the ‘Benthic Group’ of the ‘Gulf of Riga Project’. Organic carbon balance of the Gulf of Riga. In: Corner E. and Larsen B. Lipid biochemistry and reproductive biology in two species of Gammaridae.S. 1999.B. and Sandler H... Lagzdins G. Ophelia ͑Suppl. Ekebom J.. 1986.and meiofauna. Mar. 35: 193–205. Racjunas L.F.D. Mar.. inermis ͑Krøyer͒.. The Biological References Aigars J. Dybern B. Mar. Nalepa T. Frez W. 1978. Falk-Petersen S.. University of Latvia͒ for sampling and performing the excretion experiments aboard Latvian vessels.D. 46: 1714–1725.A. K. including comparisons with other areas.. Crustacea͒ species. Essink and two anonymous reviewers improved the ﬁnal version of the manuscript.G. 56: 41–48. Limnologica 15: 395–401.B. Sci.. The Baltic Marine Biologists. 1992. 1985. Seasonal changes in lipid content and composition in the benthic amphipods Monoporeia affınis and Pontoporeia femorata. Fish. Effect of bioturbation on microbial nitrogen transformations in the sediment and ﬂuxes of ammonium and nitrate to the overlying water. pp... Diurnal oxygen consumption and activity of two Pontoporeia ͑Amphipoda. New York.B. On the biology and production of Pontoporeia affınis Lindstr. Ecol.. 1993. University of Latvia͒ are thanked for delivering the benthos data. ͑eds͒. Andersin A. 1987. Monoporeia affınis ͑Lindström͒ and Pontoporeia femorata ͑Kröyer͒. J.. Seasonal course of soft-bottom community respiration in a northern Baltic archipelago. 54: 209–224.. Ivanovici A. 88: 247–263. Skadsheim A.R. Exp...L. Particle ﬂux and properties affecting the fate of bacterial productivity in the benthic boundary layer at a mud-bottom site in South-Central Gulf of Riga. 1985. Sargent J. 1984. In: Naylor E. 1986.A. and Lenshs E. SMHI Reports Oceanography No. 32: 1226–1238. Limnol. Nitrogen mineralization and denitriﬁcation in Lake Michigan sediments. Physiological procedures. 1985. Clarke A. Cavaletto J. Ecol. Floderus S. and Hartnoll R. Ecol. The Effects of Stress and Pollution on Marine Animals. Chap. and Lagzdins G.455 Hans Cederwall ͑University of Stockholm͒ and Vadims Jermakovs ͑Institute of Aquatic Ecology. 1976. USA. Proc. ͑in Russian͒ Le Borgne R.. 244: 137–44. Sedimentation of vernal and late summer phytoplankton communities in the coastal Baltic Sea. currents. Biol. Can. 101-102. Saule A. Exp. Gardner W. Bitenieks Y.. Latvian. Juris Aigars and Parsla Pallo ͑Institute of Aquatic Ecology. Can. 2002. in the Gulf of Bothnia. Sea. and Sarvala J. and Holmes L. Limnol Oceanogr 37: 1280–1289. Köuts T. Byrén L.N. 1981.. Gatten R. Livingstone D. Astrauskas A. 1978. 4͒: 17-26. 109 pp. Mar. Aukštikal’nene A. 1989. Mar. Micromethod for lipids in aquatic invertebrates. with particular reference to the relationship between macro.A. and Sandler H. Oceanogr. Lassig J. Rasmussen M. Ejdung G.. 98 pp. Dixon D. 1. and Jermakovs V. Sc. 131: 175–198.C. Comments by Dr. J. Norrköping 1990. 1983.. Cyclic Phenomena in Marine Plants and Animals.. Elmgren R. Biol. Jermakovs V. Andersin A. Pergamon Press. and Andrušaitis A. New York. Fish Aquat Sci. Ecol. Aquat. Bertil Widbom ͑University of Stockholm͒. Bull. 1992.M. Bousﬁeld E. 1996. Marine Systems 23: 233–250. 23.. Biol. Sci. J.C.R. In: Virbickas J.M. Biological Resources of Waterbodies in the Basins of the Baltic Sea. and Jensen A. Carman R. Acad. 161-178. 1999. Nutrient transport through the water-sediment interface in the Gulf of Riga.. Brown D. Growth and production of three macrozoobenthic species in the Gulf of Riga. 281: 109–121. Benthic predator-prey interactions: evidence that adult Monoporeia affınis ͑Amphipoda͒ eat postlarval Macoma balthica ͑Bivalvia͒. Vilnius. Byrén L. Elmgren R..S. Cederwall H.F. J.. Andrušaitis G. Geol. sea levels and nutrients in the Gulf of Riga. Quigley M.S. Moore M.I. Parkkonen L.M. ͑eds͒. The release of soluble end products of metabolism. and Saarso M.. B 10: 73–77. Finnish Mar. J.R. Kieler Meeresforsch Sonderh 4: 1–22. Revised morphological relationships within the amphipod genera Pontoporeia and Gammaracanthus and the ‘glacial relict’ signiﬁcance of their postglacial distributions. and Gordon W. Comparing rate and depth of feeding in benthic deposit-feeders: a test on two amphipods.. Bergström I. Carbon and nutrient geochemistry of the surface sediments of the Gulf of Riga. Proceedings of the 22nd Scientiﬁc Conference on Investigations of Baltic Waterbodies.. 134: 57–76. 2000. 1987. University of Latvia͒ and Ann-Britt Andersin ͑FIMR͒ are also thanked for most enjoyable co-operation. Ackefors H. Proceedings of the 13th European Marine Biology . Lassig J. Mar. Jähmlich S. Elmgren R. Structure and dynamics of Baltic benthos communities. Special thanks are given to Ieva Upeniece ͑Institute of Aquatic Ecology.E. 309–316. Hydrobiologia 393: 201–210. 1999. and Parrish C. 30: 1099–1105. Symposium.R... New York. Gardner W. Ecological investigations on the zooplankton community in Balsfjorden. Exp..A. 1979. J. ͑eds͒ 1995. Elmgren R. ͑eds͒. Hill C. Long-term changes in the soft-bottom macrofauna of the Gulf of Riga. Henriksen K. Priede S.J.C. 1994. and T. USA. New York. Lowe D. Bayne B. northern Norway: seasonal changes in the lipid class composition of Meganyctiphanes norvegica ͑M. Longterm ﬂuctuations of the soft bottom macrofauna in the deep areas of the Gulf of Bothnia 1954-1974. Andrušaitis A. and Widdows J. Res. Heiskanen AS and Kononen K. and Hopkins C. Observations of water exchange. and Håkansson B. Sars͒. Stebbing A. Hydrobiol. Cederwall H. Cichocki E. pp. 253: 243–251. Swedish Hydrological and Meteorological Institute Report. Recommendations on methods for marine biological studies in the Baltic Sea. Biol. O’Hara S. with special reference to Pontoporeia affınis Lindström ͑Amphipoda͒.L. The change in benthic macrofauna as an indication of eutrophication in the southern part of the Gulf of Riga.. Praeger Publishers. Ejdung G. In: Proceedings of the 17th CBO conference.. Thysanoessa raschii ͑M.
123: 555–564. Seasonal stages of phytoplankton community structure and sinking loss in the Gulf of Riga. Hietanen S. UK. 1999. Andruzaitis A. J. Oxford Science Publications. Ecol. P͒ in the pelagic system of the Gulf of Riga. Estuarine Coastal Shelf Sci. Ouellet P.P. 1999. A quantitative survey of the benthic meiofauna in the Gulf of Riga ͑eastern Baltic Sea͒. 1991.. 1996a. J. Lehtonen K. Ojaveer E. Estuarine Coastal Shelf Sci.. Parsons T. Ecol. Perspectives of research on detritus: do factors controlling the availability of detritus to macroconsumers depend on its source? J. Pelegrí S.S. Studien über die Bodentierwelt in südﬁnnländischen Küstengewässern III. 111: 237– 246. Metabolic effects of short-term starvation on the benthic amphipod Pontoporeia affınis Lindström from the northern Baltic Sea. 37: 113–127. Finnish Mar. 1999. Reigstad M. Oxford. Scient.G. Ecol. Widbom B. and their potential contribution to benthic nitrogen mineralization. Zur Morphologie und Biologie des Amphipoden Pontoporeia affınis. Cambridge University Press. Cammen L.G. Population dynamics. Taggert C. Aquat. 126: 645–654. Pallo P. Yurkovskis A. pore water proﬁles and denitriﬁcation in sediment inﬂuenced by algal sedimentation and bioturbation by Monoporeia affınis. 40: 473–480. 1991.K. Prog. and Frank K. Lipid condition and survival in shrimp Pandalus borealis larvae. Ecosystem of the Gulf of Riga between 1920 and 1990. Limnol. Biol. 7: 1–181. J. Lehtonen K. 1994. Determination of ammonia in natural waters by the phenolhypochlorite method. Ecophysiology of the benthic amphipod Monoporeia affınis in an open-sea area of the northern Baltic Sea: seasonal variations in oxygen consumption and ammonia excretion. Findlay S. Olesen M. Seasonal and spatial variation of suspended and sedimented nutrients ͑C. and Ackman R. I. Marine Syst.. Leppänen J. response to sedimentation and role in benthic metabolism of the amphipod Monoporeia affınis in an open-sea area of the northern Baltic Sea. Cycling of organic matter during the vernal growth period in the open northern Baltic proper. Fenn. Biol. 23: 211–232. Ser.M.K. 109–164. 143: 87–98. Rahm L. 1998.P. Lehtonen K. 1999. . 1969. The amphipod Pontoporeia femorata ͑Kröyer͒. Wulff F. and Heiskanen A. and Phillips N. Biol. Res. Lehtonen K.T. and Wassmann P. Nielsen L. 23: 165–184. Mäkelä K.G. 1998. with special reference to the structure of nematode assemblages. Mar.. Biological Oceanographic Processes. N. 255: 97–118. Bioturbation effects of the amphipod Corophium volutator on microbial nitrogen transformations in marine sediments. 2nd ed. J. Pelegrí S. Marine Syst. Mar. 1995. Ser. UK. Ecol. Marine Systems 23: 197–210. Peters R. ͑ed. Mar. Elmgren R. Tuominen L. 1988. and Blackburn T. Takahashi M. pp..R. Res. 49: 83–97. Denitriﬁcation in estuarine sediment stimulated by the irrigation activity of the amphipod Corophium volutator. 105: 285– 290. Baltic Sea. Prog.. Exp. Lundsgaard C. Geographical variability in the bioenergetic characteristics of Monoporeia/Pontoporeia spp. with bioenergetic considerations.. Sci. and Blackburn T.B..͒ 1995. femorata in a Baltic archipelago in relation to environmental factors. 1994. 176: 269–283. Sinking of particulate matter. 1982.K. Nutrient ﬂuxes.R. Lehtonen K.. Reigstad M. Localization of a marine source of odd chain-length fatty acids. Heiskanen A.K. Solórzano L. Mar. Mar. Ser. Pergamon Press.E. Ser.. Paradis M. 74: 99–107. Haahti H. Biol.P. Oxford.K. populations from the northern Baltic Sea. Clarendon Press.456 Chemistry of Marine Copepods. VI. 1996b. Prog. 1977. Prog. Oceanogr. Ecophysiology of the benthic amphipod Monoporeia affınis in an open-sea area of the northern Baltic Sea: seasonal variations in body composition. and Hargrave B. Comment. and Andersin A. Ophelia 49: 117–139. 277 pp. and Ólafsson E. Mar. Biol. Seasonal growth of the benthic amphipods Pontoporeia affınis and P. 121: 253–258. UK. 1937. Effects of disturbance by benthic amphipod Monoporeia affınis on meiobenthic community structure: a laboratory approach.T. Biol. 1994. Lipids 11: 863–870. Soc. Mar..K. and Kuparinen J. Cambridge.K. Estonian Academy of Sciences. J.T. 49: 368–378. The Ecological Implications of Body Size.H. Can. and RodriguezMedina M. A nutrient budget of the Gulf of Riga... Tallinn. Segerstråle S. Estonian Academy Publishers. Ecol. Mar. Fish. and Sarvala J. Uitto A. Lehtonen K. Mar.S.. 1992. 14: 799–801. nebst einer Revision der Pontoporeia-Systematik. Tenore K. Olli K. 1976. Mar. Sources of settling material: aggregation and zooplankton mediated ﬂuxes in the Gulf of Riga. 1983. 1993. 168: 71–85.H. and Olli K.H.. Ólafsson E.