Water motion is a key determinant of marine macroalgal production, influencing directly or indirectly physiological rates and community structure. Our understanding of how marine macroalgae interact with their hydrodynamic environment has increased substantially over the past 20 years, due to the application of tools such as flow visualization to aquatic vegetation, and in situ measurements of seawater velocity and turbulence. This review considers how the hydrodynamic environment in which macroalgae grow influences their ability to acquire essential resources and how macroalgae might respond physiologically to fluctuations in their hydrodynamic regime with a focus on: (1) the biochemical processes occurring within the diffusion boundary layer (DBL) that might reduce rates of macroalgal production; (2) time scales over which measurements of velocity and DBL processes should be made, discussing the likelihood of in situ mass transfer limitation; (3) if and how macroalgal morphology influences resource acquisition in slow flows; and (4) ecobiomechanics and how hydrodynamic drag might influence resource acquisition and allocation. Finally, the concept that macroalgal production is enhanced in wave-exposed versus sheltered habitats is discussed.

Marine life is controlled by multiple physical and chemical drivers and by diverse ecological processes. Many of these oceanic properties are being altered by climate change and other anthropogenic pressures. Hence, identifying the influences of multifaceted ocean change, from local to global scales, is a complex task. To guide policy-making and make projections of the future of the marine biosphere, it is essential to understand biological responses at physiological, evolutionary and ecological levels. Here, we contrast and compare different approaches to multiple driver experiments that aim to elucidate biological responses to a complex matrix of ocean

Mechanisms for inorganic carbon acquisition in macroalgal assemblages today could indicate how coastal ecosystems will respond to predicted changes in ocean chemistry due to elevated carbon dioxide (CO 2 ). We identified the proportion of noncalcifying macroalgae with particular carbon use strategies using the natural abundance of carbon isotopes and pH drift experiments in a kelp forest. We also identified all calcifying macroalgae in this system; these were the dominant component of the benthos (by % cover) at all depths and seasons while cover of noncalcareous macroalgae increased at shallower depths and during summer. All large canopy-forming macroalgae had attributes suggestive of active uptake of inorganic carbon and the presence of a CO 2 concentration mechanism (CCM). CCM species covered, on average, 15-45% of the benthos and were most common at shallow depths and during summer. There was a high level of variability in carbon isotope discrimination within CCM species, probably a result of energetic constraints on active carbon uptake in a low light environment. Over 50% of red noncalcifying species exhibited values below À30% suggesting a reliance on diffusive CO 2 uptake and no functional CCM. Non-CCM macroalgae covered on average 0-8.9% of rock surfaces and were most common in deep, low light habitats. Elevated CO 2 has the potential to influence competition between dominant coralline species (that will be negatively affected by increased CO 2 ) and noncalcareous CCM macroalgae (neutral or positive effects) and relatively rare (on a % cover basis) non-CCM species (positive effects). Responses of macroalgae to elevated CO 2 will be strongly modified by light and any responses are likely to be different at times or locations where energy constrains photosynthesis. Increased growth and competitive ability of noncalcareous macroalgae alongside negative impacts of acidification on calcifying species could have major implications for the functioning of coastal reef systems at elevated CO 2 concentrations.

In conditions of low water motion (< 0.06 m s -1), the availability of essential nutrients to macroalgae, and thus their potential productivity, may be limited by thick diffusion boundary-layers at the thallus surface. The ability of macroalgae to take up nutrients in slow moving water may be related to how their blade morphology affects diffusion boundarylayer thickness. For the giant kelp, Macrocystis integrifolia Bory, morphological measurements indicate that blades of plants from a site exposed to wave action are thick, narrow and have a heavily corrugated surface. In contrast, blades from a site with a low degree of water motion are relatively thin, with few surface corrugations and large undulations along their edges. The aim of our work was to test the hypothesis that morphological features of M. integrifolia blades from a sheltered site allow enhanced inorganic nitrogen uptake at low seawater velocities compared to blades with a wave-exposed morphology. The rate of nitrate and ammonium uptake by morphologically distinct blades of M. integrifoIia, from sites that were sheltered from and exposed to wave action, were measured in the laboratory at a range of seawater velocities (0.01 to 0.16 m s-1), between March and May 1993. For both sheltered and exposed blade morphologies, nitrate and ammonium uptake rates increased with increasing sea- water velocity, reaching a maximum rate at 0.04 to 0.06 m s-1. Uptake parameters Vmax (maximum uptake rate) and Uo.37 (the velocity at which the uptake rate is 37% of the maximum rate) were estimated using an exponential decay formula. These parameters were similar for both blade morphologies, at all seawater velocities tested. Additional measurements suggest that the nitrogen status of M. integrifolia blades from wavesheltered and exposed sites were similar throughout the experimental period, and thus nitrogen status did not affect the rate of nitrogen uptake in these experiments. On the basis of these results, we conclude that blade morphology does not enhance nitrogen uptake by M. integrifoIia in conditions of low water motion. Potential effects of diffusion boundary-layers on kelp productivity are discussed.

Coastal ecosystems that are characterized by kelp forests encounter daily pH fluctuations, driven by photosynthesis and respiration, which are larger than pH changes owing to ocean acidification (OA) projected for surface ocean waters by 2100. We investigated whether mimicry of biologically mediated diurnal shifts in pH-based for the first time on pH time-series measurements within a kelp forest-would offset or amplify the negative effects of OA on calcifiers. In a 40-day laboratory experiment, the calcifying coralline macroalga, Arthrocardia corymbosa, was exposed to two mean pH treatments (8.05 or 7.65). For each mean, two experimental pH manipulations were applied. In one treatment, pH was held constant. In the second treatment, pH was manipulated around the mean (as a step-function), 0.4 pH units higher during daylight and 0.4 units lower during darkness to approximate diurnal fluctuations in a kelp forest. In all cases, growth rates were lower at a reduced mean pH, and fluctuations in pH acted additively to further reduce growth. Photosynthesis, recruitment and elemental composition did not change with pH, but d 13 C increased at lower mean pH. Including environmental heterogeneity in experimental design will assist with a more accurate assessment of the responses of calcifiers to OA.