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Full Text in Pdf Format MARINE ECOLOGY PROGRESS SERIES Published April 25 Mar Ecol Prog Ser l Production within dense mats of the filamentous macroalga Chaetomorpha linum in relation to light and nutrient availability Dorte Krause-Jensen*,Karen McGlathery, Sarren Rysgaard, Peter Bondo Christensen National Environmental Research Institute, Vejlsevej 25, PO Box 314. DK-8600 Silkeborg, Denmark ABSTRACT. Dense mats of Chaetomorpha Iinum were incubated in the laboratory at low and high sur- face irradiance and were enriched by a simulated sediment nutrient flux. Algal activity resulted in marked diurnal variations and steep vertical gradients In O2and NH,' concentration profiles within the mats. In the light, O2 production caused supersaturation in the surface layers, and algal assimilation significantly reduced the flux of nutrients to the water column. The depth gradients of decreasing light and increasing nutrient availability within the mat were reflected in the algal tissue composition. At h~ghsurface irradiance, chlorophyll concentrations increased towards the bottom of the mat and C/N ratios gradually declined. This pattern suggested light limitation in the bottom of the mat and progres- sive N limitation towards the mat surface. Algal productivity declined with depth in the mats, reflect- ing a pronounced self-shading, and the photic zone (i.e.the depth of 1?:, surface irradiance) was only 8 cm deep. Productivity per unit volume was high, and comparisons to communities of other benthic macrophytes, benthic m~croalgae,and phytoplankton demonstrated a general pattern of increasing volume-specific productivity at decreasing extension of the photic zone, whereas the area productivity (depth-integrated)of the different plant communities is remarkably uniform. As algal density and self- shading increases, the algal mats can switch from being net productive to a status where consumption exceeds production. Reduced irradiance and increased water temperature may also trigger this shift, and the resulting effects on 0,and nutnent balances make shallow macroalgal-dominated systems inherently unstable. KEY WORDS: Macroalgal mats Chaetomorpha linum . Light . Production . Oxygen . Nutrients . Chlorophyll . Tissue composition . Microelectrodes INTRODUCTION Light and nutrients are important regulating factors of macroalgal growth and productivity. Growth can be Ephen~eralmacroalgae are often the dominant pri- described as a saturating function of both irradiance mary producers in shallow, eutrophic bays where they (Sand-Jensen & Madsen 1991) and tissue nutrient con- may form dense mats on the sediment surface (Sfriso et tent (Lavery & McComb 1991a, Pedersen 1993), but al. 1987, Lavery et al. 1991, Valiela et al. 1992). The light and nutrient effects are also strongly interactive development of algal mats can be important to the (Lapointe & Tenore 1981, Coutinho & Zingmark 1993). nutrient dynamics of coastal waters because actively The relative importance of these growth limiting fac- growing mats may intercept the nutrient flux from tors typically changes over the season. Light is the the sediment to the water column (K. McGlathery, major limiting factor during winter and early spring D. Krause-Jensen, S. Rysgaard & P. B. Christensen when nutrients are in ample supply, whereas nutrients unpubl.). usually limit production during summer when nutrient availability is low and light availability is higher (Ole- sen 1989, Pedersen 1993). In addition, spatial differ- ences between nutrient and light limitation may occur O Inter-Research 1996 Resale of fill1 article not permitted 208 Mar Ecol Prog Ser 134: 207-216, 1996 within the same estuary between sh.all.otv versus deep sites and between nutrient-poor versus nutrient-rich regions (Christensen et al. 1994). Macroalgae ca.n acclimate to these variations in light and nutrient avai.1- ability through biochemical changes to maximize pho- tosynthetic capacity and growth. For example, tissue N and pigment contents tend to increase when irradiance decreases (Markager & Sand-Jensen 1994 and refer- ences therein) and when N availability increases (Bird et al. 1982, Lapointe & Duke 1984). The relative importance of nutrients and light as growth regulating factors can also be expected to vary on a vertical scale within dense algal mats, since the microclimate can vary significantly from the surface to bottom layers of the mat (Lavery & McComb 1991b, Thybo-Christesen et al. 1993, Christensen et al. 1994). While seaweeds typically obtain nutrients from the water column, dense macroalgal mats may also benefit from sediment nutnent sources. A gradient in ambient nutrient concentrations may develop within algal mats, as the availability decreases with distance from the Fig. 1. Experimental setup. A: reservoir containing artiflciai sediment (Lavery & McComb 1991b, Thybo-Christe- seawater without nutrients; B: reservoir c0ntainin.g artificial sen et al. 1993, McGlathery et al. unpubl.). Decomposi- seawater-nutrient solution; C: peristaltic pump; D: 400 W mer- cury lamp; E: acrylic box continuously flushed with cold water tion and mineralization of organic bound nutrients in to absorb heat radiation from the lamp; F: upper compartment the bottom layers of dense algal mats may provide an containing the algal mat; G: lower compartment flushed with additional nutrient source within the mats (Thybo- nutrient solut~on;H: filter paper separating the 2 compart- Christesen & Blackburn 1993, McGlathery et al. ments, 1: teflon-coated magnets dnven by a large external unpubl.). High algal density, on the other hand, also rotating magnet; J: O2microsensor creates pronounced self-shading, which results in decreasing light availability from the surface to bottom algal mats recorded in the field (Sfriso et al. 1987, 1993, layers of the algal mats (Gordon & McComb 1989). Lavery et al. 1991, Valiela et al. 1992, Ertebjerg et al. The purpose of this work was to study macroalgal 1993, Viaroli et al. 1993). Nutrient efflux from the sed- productivity in relation to the vertical gradients of light iment was simulated by pumping a concentrated nutri- and nutrient availab~litycreated within dense algal ent solution through the lower compartment causing mats and to determine the effect of macroalgal mats on nutrients to diffuse into the algal mat across a filter oxygen and nutrient concentrations in the water col- paper separating the 2 compartments. The flux rate umn during situations of net growth and net decompo- was controlled by regulating the flow rate of nutrients sition of the algal biomass. through the lower compartment. The algae were allowed to adapt to these experimental conditions for 6 d before sampling was initiated. MATERIALS AND METHODS The upper c0mpartmen.t received a continuous sup- ply of artificial seawater Grasshoff et al. 1983) Experimental setup. An experimental setup simulat- without any nutrients added. The seawater was ing an algal matkediment system was constructed in pumped into the compartment at a flow rate of 3.2 m1 the laboratory using a 2-compartment Plexiglass cylin- min-l from an aerated reservoir to stabilize O2 and CO2 der (inner diameter 10 cm, height 40 cm; Fig. 1). The concentrations. The water column above the algal mat filamentous macroalga Chaetomorpha linum was col- was stirred by a magnetic stirrer bar (60 rpm) posi- lected in the shallow cove Kertinge Nor, Denmark, and tioned 5 cm above the mat surface. added to the upper compartment to create a 15 cm Nutrients were added to artificial seawater in the thick algal mat comparable to in situ conditions. The lower compartment to reach a final concentration of algal mat consisted of 30 g (fresh weight, fw) actively 650 pM NH,' and 65 pM Pod3-.The nutrient solution growing algae overlying 5 g fw of black algal filaments was aerated with N2 to make the water anoxic before it taken from the bottom of an algal mat. This density was pumped through the lower compartment. Several corresponds to 4.5 kg fw m-2 (or approximately 600 g tests of the outflowing water verified that bacterial dw m 7,which resembles the maximum biomass of growth (e.g. of nitrifying bacteria) in the lower cham- Krause-Jensen et al.. Light and nutrient availability in algal mats ber was negligible. The nutrient concentrations and ity (P)of each 2 cm section of the mats was calc.ulated flow rate were chosen to obtain an efflux of approxi- from: mately 1000 pm01 N m-2 h-' and 100 pm01 P m-, h-', P (pm01 C g dw-' h-') = corresponding to the maximum in situ flux rates during (j3C%end- 13C%start)X TC X ('2D~C/'3DIC)/t summer from the sediments in Kertinge Nor (Chris- (2) tensen et al. 1994). The flow rate (f), and the concen- where I3C%start and j3C%end represent the 13C con- trations of nutrients in the inflow (Ci)and the outflow tent of Chaetomorpha linum tissue before and after (C,,) were measured every 3 to 5 h during the experi- incubation with NaHI3CO3,respectively. TCis the total ment, and the actual flux rate (F)was calculated as: C content, 12DIC/'3~~Cthe average specific activity of DIC (disolved inorganic carbon) during the incubation, F = (C,- C,) X f/A (1) and t the incubation time. where A is the surface area separating the 2 chambers. The 13C content of DIC in the incubation water was Light was supplied from a 400 W mercury lamp in a determined in 2 m1 water samples collected through 12 h light/l2 h dark cycle. One core received approxi- silicone stoppered ports in the cylinder wall. The sanl- mately 120 pm01 photons m-, S-' (referred to as 'low ples were stored in 1.5 m1 borosilicate glass vials with light') while the other core received approximately 380 PTFE-coated Butyl rubber stoppers, and bacterial pm01 m-2 ss1 ('high light'). The cores were wrapped in activity was stopped by adding 20 p1 20 % ZnCl before black plastic to prevent light exposure through the the samples were transferred to helium filled 5 m1 glass sides. The experiments were run in a temperature- vials (Exetainer, Labco, UK).
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