Scientia Marina, Vol 75, No 4 (2011)

Photosynthesis and daily metabolic carbon balance of the invasive Caulerpa racemosa var. cylindracea (Chlorophyta: Caulerpales) along a depth gradient

Jaime Bernardeau-Esteller
Instituto Español de Oceanografía, Centro Oceanográfico de Murcia - Departamento de Ciencias del Mar y Biología Aplicada, Universidad de Alicante , Spain

Lázaro Marín-Guirao
Instituto Español de Oceanografía, Centro Oceanográfico de Murcia , Spain

Jos M. Sandoval-Gil
Instituto Español de Oceanografía, Centro Oceanográfico de Murcia - Departamento de Ciencias del Mar y Biología Aplicada, Universidad de Alicante , Spain

Juan M. Ruiz
Instituto Español de Oceanografía, Centro Oceanográfico de Murcia , Spain


The photosynthetic plasticity of the invasive green alga Caulerpa racemosa v. cylindracea (hereafter C. racemosa) has been proposed as a relevant mechanism determining its successful performance on Mediterranean benthic assemblages over broad depth gradients. In the present study, the photosynthetic performance of C. racemosa was evaluated through a carbon balance approach at three invaded sites with contrasting depths (11, 18 and 26 m) and light regimes. At each sampling depth, photosynthesis vs irradiance (P vs E) curves were performed on C. racemosa fronds and daily net productivity values were obtained by the numerical integration of P vs E models with continuous recording of irradiance measured on the sea floor. Photosynthetic responses were consistent with those typically exhibited by shade-adapted macroalgal species and other Mediterranean populations of C. racemosa: a significant reduction in maximum photosynthesis (Pmax) occurring at an intermediate depth (18 m) and a higher photosynthetic efficiency (α) and lower dark respiration rate (Rd) at the deepest sampling depth. Mean values of daily net C balance obtained from the deeper site were only 15% lower than those obtained from the shallower site, despite the severe reduction in light availability. This daily net carbon gain was ca. 29% higher than would be expected if photosynthetic adjustments did not occur in the deeper algal population. The evidence provided by these data support the hypothesis of photoacclimation in C. racemosa as an effective mechanism to optimize algal productivity across depth gradients in the Mediterranean Sea.


Caulerpa racemosa; carbon balance; invasive species; photosynthesis; depth; Mediterranean

Full Text:



Argyrou, M., A. Demetropoulos and M. Hadjichristophorou. – 1999. Expansion of the macroalga Caulerpa racemosa and changes in softbottom macrofaunal assemblages in Moni Bay, Cyprus. Oceanol. Acta, 22: 517-528.

Balata, D., L. Piazzi and F. Cinelli. – 2004. A comparison among assemblages in areas invaded by Caulerpa taxifolia and Caulerpa racemosa on a subtidal Mediterranean rocky bottom. Mar. Ecol., 25(1): 1-13.

Baly, E.C.C. – 1935. The kinetics of photosynthesis. Proc. R. Soc. Lond. Ser. B Biol. Sci., 117: 218-239.

Boudouresque, C.F. and M. Verlaque . – 2002. Biological pollution in the Mediterranean Sea: invasive versus introduced macrophytes. Mar. Poll. Bull., 44: 32-38.

Bulleri, F. and L. Benedetti-Cecchi. – 2008. Facilitation of the introduced green alga Caulerpa racemosa by resident algal turfs: experimental evaluation of underlying mechanisms. Mar. Ecol. Prog. Ser., 364: 77-86.

Cayabyab, N.M. and S. Enríquez. – 2007. Leaf photoacclimatory responses of the tropical seagrass Thalassia testudinum under mesocosm conditions: a mechanistic scaling-up study. New Phytol., 176: 108-123. PMid:17696981

Cebrian, E. and E. Ballesteros. – 2009. Temporal and spatial variability in shallow and deep water populations of the invasive Caulerpa racemosa var. cylindracea in the western Mediterranean. Estuar. Coast. Shelf Sci., 83: 469-474.

Cecherelli, G. and D. Campo. – 2002. Different effects of Caulerpa racemosa on two co-ocurring seagrasses in the Mediterranean. Bot. Mar., 45: 71-76.

Chisholm, J.R.M. and J.M. Jaubert. – 1997. Photoautotrophic metabolism of Caulerpa taxifolia (Chlorophyta) in the NW Mediterranean. Mar. Ecol. Prog. Ser., 153: 113-123.

Dunton, K.H. and D.M. Shell. – 1986. Seasonal carbon budget and growth of Laminaria solidungula in the Alaskan High Artic. Mar. Ecol. Prog. Ser., 31: 57-66.

Falkowski, P.G. and J.A. Raven. – 1997. Aquatic photosynthesis. Blackwell Scientific Publishers, Oxford.

Flagella, M.M., M. Lorenti and M.C. Buia. – 2008. Temperature response in a shallow-water Mediterranean population of Caulerpa racemosa var. cylindracea (Caulerpales, Chloropyta), and a possible strategy of season anticipation. Bot. Mar., 51(4): 278-284.

Gacia, E., C. Rodríguez-Prieto, O. Delgado and E. Ballesteros. – 1996. Seasonal light and temperature responses of Caulerpa taxifolia from northwestern Mediterranean. Aquat. Bot., 53: 215-225.

Gatusso, J.P. and J. Jaubert. – 1985. Photosynthesis and respiration of Caulerpa racemosa (Chlorophyceae, Caulerpales) grown in aquaria: Effects of light and temperature. Bot. Mar., 28: 327-332.

Gómez, I., G. Weykam, H. Klöser and C. Wiencke. – 1997. Photosynthetic light requirements, metabolic carbon balance and zonation of sublittoral macroalgae from King George Island (Antarctica). Mar. Ecol. Prog. Ser., 148: 281-293.

Gómez, I. – 2001. Ecophysiology of antartic macroalgae: effects of environmental light conditions on photosynthetic metabolism. Rev. Chil. Hist. Nat., 74: 251-271.

Johansson, G. and P. Snoeijs. – 2002. Macroalgal photosynthetic responses to light in relation to thallus morphology and depth zonation. Mar. Ecol. Prog. Ser., 244: 63-72.

Kirk, J.T. – 1994. Light and Photosynthesis in Aquatic Ecosystems. Cambrige University Press. Cambridge.

Klein, J. and M. Verlaque. – 2008. The Caulerpa racemosa invasion. A critical review. Mar. Poll. Bull., 56: 205-225. PMid:18061625

Klein, J.and M. Verlaque. – 2009. Macroalgal assemblages of disturbed coastal detritic bottoms subject to invasive species. Estuar. Coast. Shelf Sci., 82: 461-468.

Littler, M.M., D.S. Littler, S.M. Blair and J.N. Norris. – 1986. Deep-water plant communities from an unchartered seamount off San Salvador Island, Bahamas: distribution, abundance and primary productivity. Deep-Sea Res., I 33: 881-892.

Lobban, C.S. and P.J. Harrison. – 1997. Seaweed ecology and physiology. Cambridge University Press, Cambridge.

Matta, J.L. and D.J.Chapman. – 1991. Photosynthetic responses and daily carbon balance of Colpomenia peregrina: seasonal variations and differences between intertidal and subtidal populations. Mar. Biol., 108: 303-313.

Markager, S. and K. Sand-Jensen. – 1992. Light requirements and depth zonation of marine macroalgae. Mar. Ecol. Prog. Ser., 88: 83-92.

Markager, S. and K. Sand-Jensen. – 1994. The physiology and ecology of light-growth relationships in macroalgae. In: F.E. Round and D.J. Chapman (eds.), Progress in Phycological Research, pp. 209-298. Biopress, Bristol.

Ohba, H., H. Nashima and S. Enomoto. – 1992. Culture studies on Caulerpa (Caulerpales, Chlorophyceae) III. Reproduction, development and morphological variation of laboratory-cultured C. racemosa var. peltata. Bot. Mag. Tok., 105: 589-600.

Pérez-Lloréns, J.L., J.J. Vergara, R.R. Pino, I. Hernández, G. Peralta and F.X. Niell. – 1996. The effect of photoacclimation on the photosynthetic physiology of Ulva curvata and Ulva rotundata (Ulvales, Chlorophyta). Eur. J. Phycol., 31: 349-359.

Peterson, R.D. – 1972. Effects of light intensity on the morphology and productivity of Caulerpa racemosa (Forskål) J. Agardh. Micronesica, 8: 63-86.

Piazzi, L., G. Cecherelli. and F. Cinelli. – 2001. Threat to macroalgal diversity: effects of the introduced green alga Caulerpa racemosa in the Mediterranean. Mar. Ecol. Prog. Ser., 210: 149-159.

Piazzi, L., A. Meinesz and M. Verlaque. – 2005. Invasion of Caulerpa racemosa var. cylindracea (Caulerpales, Chlorophyta) in the Mediterranean Sea: an assessment of the spread. Cryptog. Algol., 26: 189-202.

Piazzi, L. and D. Balata. – 2008. The spread of Caulerpa racemosa var. cylindracea in the Mediterranean Sea: An example of how biological invasions can influence beta diversity. Mar. Environ. Res., 65: 50-61. PMid:17904216

Quinn, G.P. and M.J. Keough. – 2002. Experimental designs and data analysis for biologists. Cambridge University Press, Cambridge.

Ramus, J., F. Lemons and C. Zimmerman. – 1977. Adaptation of light-harvesting pigments to downwelling light and the consequent photosynthetic performance of the eulittoral rockweeds Ascophyllum nodosum and Fucus vesiculosus . Mar. Biol., 42(4): 293-303.

Raniello, R., M. Lorenti, C. Brunet and M.C. Buia. – 2004. Photosynthetic plasticity of an invasive variety of Caulerpa racemosa in a coastal Mediterranean area: light harvesting capacity and seasonal acclimatation. Mar. Ecol. Prog. Ser., 271: 113-120.

Raniello, R., M. Lorenti, C. Brunet and M.C. Buia. – 2006. Photoacclimation of the invasive alga Caulerpa racemosa var. cylindracea to depth and daylight patterns and a putative new role for siphonaxanthin. Mar. Ecol., 27: 20-30.

Riechert, R. and C.J. Dawes. – 1986. Acclimation of the green alga Caulerpa racemosa var. uvifera to light. Bot. Mar., 29: 533-537.

Robledo, D. and Y. Freile-Pelegrín. – 2005. Seasonal variations in photosynthesis and biochemical composition of Caulerpa spp (Bryopsidales, Chlorophyta) from the Gulf of Mexico. Phycologia, 44: 312-319.[312:SVIPAB]2.0.CO;2

Rosenberg, G., D.S. Littler, M.M. Littler and C. Oliveira. – 1995. Primary production and photosynthetic quotients of seaweeds from Sao Paulo State, Brazil. Bot. Mar., 38: 369-377.

Ruitton, S., M. Verlaque and C.F. Boudouresque. – 2005. Seasonal changes of the introduced Caulerpa racemosa var. cylindracea (Caulerpales, Chlorophyta) at the northwest limit of its Mediterranean range. Aquat. Bot., 82: 55-70.

Ruitton, S., M. Verlaque, G. Aubin and C.F. Boudouresque. – 2006. Grazing on Caulerpa racemosa var. cylindracea (Caulerpales, Chlorophyta) in the Mediterranean Sea by herbivorous fishes and sea urchins. Vie Milieu, 56: 33-41.

Ruiz, J.M. and J. Romero. – 2001. Effects of in situ experimental shading on the Mediterranean seagrass Posidonia oceanica. Mar. Ecol. Prog. Ser., 215: 107-120.

Ruiz, J.M., L. Marín-Guirao, J. Bernardeau-Esteller, A. Ramos-Segura, R. García-Muñoz and J.M. Sandoval-Gil. – 2011. Spread of the invasive alga Caulerpa racemosa var. cylindracea (Caulerpales, Chlorophyta) along the Mediterranean coast of the Murcia Region (SE Spain). Anim. Bio. Cons., 34: 73-82.

Terrados, J. and J.D. Ros. – 1992. The influence of temperature on seasonal variation of Caulerpa prolifera (Forsskal) Lamouroux photosynthesis and respiration. J. Exp. Mar. Biol. Ecol., 162: 199-212.

Tomas, F., E. Cebrian and E. Ballesteros. – 2011. Differential herbivory of invasive algae by native fish in the Mediterranean Sea. Estuar. Coast. Shelf Sci., 92: 27-34.

Streftaris, N. and A. Zenetos. – 2006. Alien Marine Species in the Mediterranean – the 100 ‘Worst Invasives’ and their impact. Medit. Mar. Sci., 7(1): 87-118.

Vázquez-Luis, M., P. Sánchez-Jerez and J.T. Bayle-Sempere. – 2008. Changes in amphipod (Crustacea) assemblages associated with shallow-water algal habitats invaded by Caulerpa racemosa var. cylindracea in the western Mediterranean Sea. Mar. Environ. Res., 65: 416-426 PMid:18342365

Verlaque, M., C. Durand, J.M. Huisman, C.F. Boudouresque and Y. Le Parco. – 2003. On the identity and origin of the Mediterranean invasive Caulerpa racemosa (Caulerpales, Chlorophyta). Eur. J. Phycol., 38: 325-339.

Walker, D.A. – 1985. Measurement of oxygen and chlorophyll fluorescence. In: J. Coombs, D.O. Hall, S.P. Long and J.M.O. Scurlock (eds.), Techniques in bioproductivity and photosynthesis. Pergamon Press, Oxford.

Williams, S.L. and W.C. Dennison. – 1990. Light availability and diurnal growth of a green macroalga (Caulerpa cupressoides) and a seagrass (Halophila decipiens). Mar. Biol., 196: 437-443.

Yokohama, Y. and T. Misonou. – 1980. Chlorophyll a:b ratios in marine benthic algae. Jap. J. Phycol., 28: 219-223.

Zimmerman, R.C., A. Capello-Pasini and R.S. Alberte. – 1994. Modeling daily production of aquatic macrophytes from irradiance measurements: a comparative analysis. Mar. Ecol. Prog. Ser., 114: 185-196.

Copyright (c) 2011 Consejo Superior de Investigaciones Científicas (CSIC)

Creative Commons License
This work is licensed under a Creative Commons Attribution 4.0 International License.

Contact us

Technical support