Progress in the oceanographic knowledge of Chilean interior waters, from Puerto Montt to Cape Horn. N. Silva & S. Palma (eds.). 2008 Comité Oceanográfico Nacional - Pontificia Universidad Católica de Valparaíso, Valparaíso, pp. 93-97. 6.2 Primary productivity and phytoplankton size and biomass in the austral Chilean channels and fjords: spring-summer patterns Vivian Montecino1 & Gemita Pizarro2 1 Facultad de Ciencias. Universidad de Chile E-mail: [email protected] 2 Instituto de Fomento Pesquero E-mail: [email protected] Phytoplankton occurs in a wide range of sizes and forms. Smaller organisms (< 5 µm) are more frequent and abundant in less productive systems, whereas larger organisms (> 20 µm), or microphytoplankton, prevail in eutrophic waters, which are more productive, being rich in phosphorus and nitrogen. Consequently, the dynamics of phytoplankton in relation to the local environment and other organisms are particularly important when the ecosystem's biological productivity is estimated. As a component of biogeochemical processes, primary productivity, on average 1 g·m–2·d–1, helps explain the function of phytoplankton in the carbon pump that reduces atmospheric CO 2. Primary productivity is studied according to the timescale on which phytoplankton photosynthetic processes and growth occur. On a smaller scale, these experiments are performed in situ or in vitro, lasting from minutes-hours or hours-days, depending on the proposed objectives. Larger scale questions deal with seasonal, intraseasonal, and interannual variability, both in the water column (vertical) and horizontally (mesoscale). In the first case, the uncertainties are physiological, 76° 75° 73°W 74° 47°S G. de Penas 92 Pacific Ocean Oceanic primary productivity, generated mainly by the microscopic autotrophic organisms that make up the phytoplankton, is an essential element in marine ecosystems. Primary productivity is a two-stage process, consisting of photosynthesis and biosynthesis. Photosynthesis, or carbon fixation, is driven by the chlorophyll contained in microalgal chloroplasts (Kirk, 1994). Chlorophyll-a concentrations (Chl-a) are universally used as a measure of phytoplankton biomass. Microalgae form associations that interact with other microorganisms, constituting a microbial web that regulates in situ nutrient and carbon recycling, its transfer to higher trophic levels, or its sedimentation to deeper waters. C. Fallos E. Steffen 91 90 5 12 7 17 E. Mitchel 48° C. Baker S. Iceberg 20 C. Messier A. Inglesa 87 28 C. Ladrillero S. Eyre 24 27 29 86 77 26 85 C. Picton E. Falcon 78 31 25 84 32 33 79 C. Trinidad S. Penguin 81 35 82 36 S. Europa 38 39 73 E. Peel 76 74 E. Calvo 42 71 70 E. Amalia 43 75 C. Concepción 88 49° 50° 51° 44 45 C. Smyth 67 48 50 Str ait of CIMAR 2 Fiordos Ma gel lan 52° 61 53° Figure 1: Geographic position of the sampling stations used to determine primary productivity and phytoplankton biomass in the CIMAR 2 Fiordos cruise. whereas, in the second case, they are ecological (Marra, 2002). The CIMAR 2-4 Fiordos cruises, carried out from Boca del Guafo to Cape Horn (Fig. 1, 2), covered the vast geographical area of austral Chilean channels and fjords, which are characterized by different water masses (Silva et al., 1998; Guzmán & Silva, 2002; Valdenegro & Silva, 2003). Here, phytoplankton biomass, — 93 — Montecino, V. & G. Pizarro P. Dungeness 27 56 54 S . til I nú B.C. Whiteside Alm 43ºS 4 Pacific Ocean ellan 7 54° a 2 Angostura 6 n ea Strait of Ma g 25 21 Oc C. Deseado S. Boca delGuafo a 1 Angostura tic an 11 20 Atl 53° ay Otw I.hC ilo é 52°S 6 C. 55° 32 urn 34 35 ckb o C C. o r e llen Ba C. 51 37 nta z go 40 41 C. Beagle 42 I. Navarino 47 B. Nassau 56° 44 I. W olla Cape Horn ston 7 Laguna San Rafael 26 27 CIMAR 3 Fiordos (Phase 2) 74° 72° 66°W 75º 68° 74° 45º 46º 28 Golfo Elefantes CIMAR 4 Fiordos 70° 44º Canal Jacaf 10 Canal Puyuguapi 12A Isla Meninea 12 13 17A 16 14 15 21Fiordo Aysén 1718 19 21A 34 22 33 23 31 Estero Quitralco 32 24 30 29 25 Estero Cupquelán 48 Pacific Ocean Canal Moraleda 8 9 ira rd wa Fro 5 73º 47º 72°W Figure 2: Geographic position of sampling stations to determine primary productivity and phytoplankton biomass in the CIMAR 3 (Phase 2) and 4 Fiordos (Phase 1 and 2) cruises. expressed as Chl-a, was measured along with species diversity (estimated through the ShannonWeaver index, H') and the variety of sizes found in the surface samples taken from the euphotic or well-lit zone. Carbon fixation was also estimated by using an incubator with an artificial light source, according to the methodology described by Pizarro et al. (2000). In the study area, the fractioning of the total biomass showed that phytoplanktonic organisms larger than 20 µm (microphytoplankton) were recurrent on meso and macro scales. Moreover, species richness was 17-27 for the maximum H' diversity values and 5-10 for the minimum H' values. The surface abundances and most recurring (> 45 %) microphytoplankton species in the three studied zones were Skeletonema costatum (67 %) in October 1998 and Guinardia delicatula (65 %) in February 1999 between Boca del Guafo and Laguna San Rafael (northern zone); Thalassiosira minuscula (91 %) in August 1995 and Chaetoceros cinctus (36 %) in October 1996 from Golfo de Penas to Strait of Magellan (central zone); and Chaetoceros sp. (56 %) in October 1998 from Strait of Magellan to Cape Horn (southern zone). The most frequent distribution pattern showed that the numerically predominant species were the same in only a few places, whereas the rarest species were found at nearly all the sites. A similar situation was observed in terms of biomass, with a heterogeneous distribution of satellite chlorophyll (Chl-sat) and high concentrations (> 10 mg Chla·m-3) at specific sites (Fig. 3). The vertical distribution of phytoplankton biomass showed a significant relationship between Chl-a at the surface (0-5 m) and at 10 m depth (Fig. 4). Sites with differences of one order of magnitude between these two depths were relatively rare. Most phytoplankton concentrations greater than 1 mg·m-3 were made up by the fraction exceeding 20 µm (Fig. 5). When considering the average vertical Chl-a profiles in the three zones, the northern zone clearly had higher and deeper concentrations (> 2 mg·m–3; > 20 m), whereas the largest abundances were found to 10 m depth in the central zone; the southern zone presented a more uniform depth distribution (≤1 mg·m–3) (Fig. 6). Comparatively, heterogeneity in the values at 20 m depth was large, and the variability was not lower at the vertical peaks, which is consistent with the classical patterns determined for estuaries. According to these patterns, the euphotic zone — 94 — Primary productivity and phytoplankton size and biomass in the austral Chilean channels and fjords: spring-summer patterns Northern, Central and Southern Zones (August 1995 - October 1998) –3 Chlorophyll-a 10 m (mg·m ) 100 10 1 0 0 1 10 100 –3 Surface Chlorophyll-a (mg·m ) Figure 4: Comparison of the chlorophyll-a values at the surface and at 10 m depth for each oceanographic station. Most of the results fit a homogeneous vertical distribution of the values. a reached an average of 20 ± 8 m depth, considering all the measurements carried out in the fjords, channels, and oceanic areas. Csat: Chlorophyll images SeaWiFs (±DS) 15 09-jan-99 -3 Csat (mg·m ) 23-jan-99 24-jan-99 10 10-mar-99 5 b 0 43 44 45 46 47 Lat S Figure 3: a) Distribution of surface chlorophyll concentrations between Puerto Montt and the Península Taitao according to the SeaWiFs satellite image of 24 March 1999 (Laboratorio de Modelación Ecológica, Universidad de Chile); b) Surface chlorophyll values between 43º and 46º S extracted from four SeaWiFs satellite images taken between January and March 1999). By graphing the average primary productivity values for each cruise and the values obtained at each one of the studied stations, spatial variability was observed to be high in the three geographic zones: average values were around 3 in the northern zone, < 1 g·m–2·d–1 in the central zone (Fig. 7), and more heterogeneous in the southern zone. The physical factors that control estuarine systems (Garret & Marra, 2002) are responsible for this variability and are consistent with the primary productivity results obtained in the three analyzed areas. Thus, vertical distribution patterns of phytoplanktonic biomass can be attributed to local differences in the intensity of the mixing and stratification processes and, therefore, to the photo-acclimatization processes of the autotrophic organisms. Similarities among the three zones indicate that the Chl-a abundance is determined by the size structure of the phytoplanktonic organisms (Montecino, 2001). — 95 — Montecino, V. & G. Pizarro 100 Chl-a fraction >20 m (mg·m ) y = 0,8962 x -0,8538 R2 = 0,9407 Total Chlorophyll (mg·m–3) 1:1 -3 0 1 2 3 4 6 5 7 0 10 Depth (m) 10 1 20 30 40 CIMAR 2 Fiordos, Central Zone 95-96 50 0 0 1 10 CIMAR 3 Fiordos, Southern Zone 98 CIMAR 4 Fiordos, Northern Zone 98-99 60 100 –3 Non fractionated Chl-a (mg·m ) Northern Zone 98 12000 Figure 6: Comparison of vertical profiles of average total chlorophyll for the northern (1998-1999), central (1995-1996), and southern (1998) zones. Northern Zone 99 Central Zone 96 Southern Zone 98 –2 –1 Daily primary production (mg·m ·d ) Figure 5: Relationship between total biomass (unfractionated chlorophyll-a) and chlorophyll-a estimated for the largest size fraction (> 20 µm) at the stations analyzed in the channel and fjord region. 10000 8000 6000 4000 2000 Aver. Northern Zone 98 Aver. Northern Zone 99 Aver. Central Zone 96 54 37 21 7 50 35 26 24 20 17 16 11Y 10 7 27 17 0 11 Station Number Aver. Southern Zone 98 Figure 7: Daily primary productivity measured in vitro by the 14C method at different stations and on different cruises from north to south. Non-shaded bars indicate the average value of primary productivity (mg·m–2·d–1) estimated for each cruise. — 96 — Primary productivity and phytoplankton size and biomass in the austral Chilean channels and fjords: spring-summer patterns Primary productivity and Chl-a abundance patterns are also congruent with studies on the quantity of organic matter in sediments and the effect of glaciers in the zone (Silva et al., 1998). These glaciers release inorganic matter known as glacial silt that, in some sectors, “dilutes” the organic content of the sediments and attenuates primary productivity due to decreased light penetration. However, photosynthetic pigments absorb light, also causing an endogenous light-limitation (Pizarro et al., 2005). This light limitation, together with the scarcity of other resources such as dissolved nutrients, tends to favor smaller-sized phytoplankton fractions as the predominant component in the total biomass. The patterns described are a tool for quantifying the variability of these ecosystems on meso and macro scales. References Garrett, A. & J. Marra. 2002. Effects of upper ocean physical processes (turbulence, advection and airsea interaction) on oceanic primary production. In: A. Robinson, J. Mc Carthy & B. Rotschild (eds.). The Sea, 12: 19-49. Guzmán, D. & N. Silva. 2002. Caracterización física y química y masas de agua en los canales australes de Chile entre boca del Guafo y golfo Elefantes (Crucero CIMAR 4 Fiordos). Cienc. Tecnol. Mar, 25(2): 45-76. freshwater ecosystems. Blackwell Science, New York, pp. 78-108. Montecino, V. 2001. Alometría y biodiversidad en fitoplancton en relación con la productividad primaria en ecosistemas pelágicos. In: K. Alveal & T. Antezana (eds.). Sustentabilidad de la biodiversidad, un problema actual, bases científicotécnicas, teorizaciones y proyecciones. Universidad de Concepción, Concepción, pp. 199-215. Pizarro, G., J. L. Iriarte, V. Montecino, J. L. Blanco & L. Guzmán. 2000. Distribución de la biomasa fitoplanctónica y productividad primaria máxima de fiordos y canales australes (47°-50° S) en octubre 1996. Cienc. Tecnol. Mar, 23: 25-48. Pizarro, G., V. Montecino, L. Guzmán, V. Muñoz, V. Chacón, H. Pacheco, M. Frangópulos, L. Retamal & C. Alarcón 2005. Patrones locales recurrentes del fitoplancton en fiordos y canales australes (46º-56º S) en primavera y verano. Cienc. Tecnol. Mar, 28(2): 63-83. Silva, N., J. Maturana, J. Sepúlveda & R. Ahumada. 1998. Materia orgánica, C y N, su distribución y estequiometría, en sedimentos superficiales de la región norte de los fiordos y canales australes de Chile (Crucero CIMAR-Fiordo 1). Cienc. Tecnol. Mar, 21: 49-74. Valdenegro, A. & N. Silva. 2003. Caracterización física y química de la zona de canales y fiordos australes de Chile entre el Estrecho de Magallanes y Cabo de Hornos (CIMAR 3 Fiordos). Cienc. Tecnol. Mar, 26(2): 19-60. Kirk, J. T. O. 1994. Light and photosynthesis in aquatic ecosystems. Cambridge University Press, London, 509 pp. Marra, J. 2002. Approaches to the measurements of plankton production. In: P. J. le B. Williams, D. N. Thomas & C. S. Reynolds (eds.). Phytoplankton productivity: carbon assimilation in marine and — 97 —
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