Sea surface microlayer
The sea surface microlayer is the boundary interface between the atmosphere and ocean, covering about 70% of Earth's surface. With an operationally defined thickness between 1 and, the SML has physicochemical and biological properties that are measurably distinct from underlying waters. Recent studies now indicate that the SML covers the ocean to a significant extent, and evidence shows that it is an aggregate-enriched biofilm environment with distinct microbial communities. Because of its unique position at the air-sea interface, the SML is central to a range of global marine biogeochemical and climate-related processes.
The sea surface microlayer is the boundary layer where all exchange occurs between the atmosphere and the ocean. The chemical, physical, and biological properties of the SML differ greatly from the sub-surface water just a few centimeters beneath.
Despite the huge extent of the ocean's surface, until now relatively little attention has been paid to the sea surface microlayer as the ultimate interface where heat, momentum and mass exchange between the ocean and the atmosphere takes place. Via the SML, large-scale environmental changes in the ocean such as warming, acidification, deoxygenation, and eutrophication potentially influence cloud formation, precipitation, and the global radiation balance. Due to the deep connectivity between biological, chemical, and physical processes, studies of the SML may reveal multiple sensitivities to global and regional changes.
Understanding the processes at the ocean's surface, in particular involving the SML as an important and determinant interface, could provide an essential contribution to the reduction of uncertainties regarding ocean-climate feedbacks. As of 2017, processes occurring within the SML, as well as the associated rates of material exchange through the SML, remained poorly understood and were rarely represented in marine and atmospheric numerical models.
Overview
The sea surface microlayer is the boundary interface between the atmosphere and ocean, covering about 70% of the Earth's surface. The SML has physicochemical and biological properties that are measurably distinct from underlying waters. Because of its unique position at the air-sea interface, the SML is central to a range of global biogeochemical and climate-related processes. Although known for the last six decades, the SML often has remained in a distinct research niche, primarily as it was not thought to exist under typical oceanic conditions. Recent studies now indicate that the SML covers the ocean to a significant extent, highlighting its global relevance as the boundary layer linking two major components of the Earth system – the ocean and the atmosphere.In 1983, Sieburth hypothesised that the SML was a hydrated gel-like layer formed by a complex mixture of carbohydrates, proteins, and lipids. In recent years, his hypothesis has been confirmed, and scientific evidence indicates that the SML is an aggregate-enriched biofilm environment with distinct microbial communities. In 1999 Ellison et al. estimated that 200 Tg C yr−1 accumulates in the SML, similar to sedimentation rates of carbon to the ocean's seabed, though the accumulated carbon in the SML probably has a very short residence time. Although the total volume of the microlayer is very small compared to the ocean's volume, Carlson suggested in his seminal 1993 paper that unique interfacial reactions may occur in the SML that may not occur in the underlying water or at a much slower rate there. He therefore hypothesised that the SML plays an important role in the diagenesis of carbon in the upper ocean. Biofilm-like properties and highest possible exposure to solar radiation leads to an intuitive assumption that the SML is a biochemical microreactor.
Historically, the SML has been summarized as being a microhabitat composed of several layers distinguished by their ecological, chemical and physical properties with an operational total thickness of between 1 and 1000 μm. In 2005 Hunter defined the SML as a "microscopic portion of the surface ocean which is in contact with the atmosphere and which may have physical, chemical or biological properties that are measurably different from those of adjacent sub-surface waters". He avoids a definite range of thickness as it depends strongly on the feature of interest. A thickness of 60 μm has been measured based on sudden changes of the pH, and could be meaningfully used for studying the physicochemical properties of the SML. At such thickness, the SML represents a laminar layer, free of turbulence, and greatly affecting the exchange of gases between the ocean and atmosphere. As a habitat for neuston, the thickness of the SML in some ways depends on the organism or ecological feature of interest. In 2005, Zaitsev described the SML and associated near-surface layer as an incubator or nursery for eggs and larvae for a wide range of aquatic organisms.
Hunter's definition includes all interlinked layers from the laminar layer to the nursery without explicit reference to defined depths. In 2017, Wurl et al. proposed Hunter's definition be validated with a redeveloped SML paradigm that includes its global presence, biofilm-like properties and role as a nursery. The new paradigm pushes the SML into a new and wider context relevant to many ocean and climate sciences.
According to Wurl et al., the SML can never be devoid of organics due to the abundance of surface-active substances in the upper ocean and the phenomenon of surface tension at air-liquid interfaces. The SML is analogous to the thermal boundary layer, and remote sensing of the sea surface temperature shows ubiquitous anomalies between the sea surface skin and bulk temperature. Even so, the differences in both are driven by different processes. Enrichment, defined as concentration ratios of an analyte in the SML to the underlying bulk water, has been used for decades as evidence for the existence of the SML. Consequently, depletions of organics in the SML are debatable; however, the question of enrichment or depletion is likely to be a function of the thickness of the SML constitutes the uppermost layer of the ocean, only 1–1000 μm thick, with unique chemical and biological properties that distinguish it from the underlying water. Due to the location at the air-sea interface, the SML can influence exchange processes across this boundary layer, such as air-sea gas exchange and the formation of sea spray aerosols.
Due to its exclusive position between the atmosphere and the hydrosphere and by spanning about 70% of the Earth's surface, the sea-surface microlayer is regarded as a fundamental component in air–sea exchange processes and in biogeochemical cycling. Although having a minor thickness of <1000 μm, the elusive SML is long known for its distinct physicochemical characteristics compared to the underlying water, e.g., by featuring the accumulation of dissolved and particulate organic matter, transparent exopolymer particles, and surface-active molecules. Therefore, the SML is a gelatinous biofilm, maintaining physical stability through surface tension forces. It also forms a vast habitat for different organisms, collectively termed as neuston with a recent global estimate of 2 × 1023 microbial cells for the sea-SML.
Life at air–water interfaces has never been considered easy, mainly because of the harsh environmental conditions that influence the SML. However, high abundances of microorganisms, especially of bacteria and picophytoplankton, accumulating in the SML compared to the underlying water were frequently reported, accompanied by a predominant heterotrophic activity. This is because primary production at the immediate air–water interface is often hindered by photoinhibition. However, some exceptions of photosynthetic organisms, e.g., Trichodesmium, Synechococcus, or Sargassum, show more tolerance towards high light intensities and, hence, can become enriched in the SML. Previous research has provided evidence that neustonic organisms can cope with wind and wave energy, solar and ultraviolet radiation, fluctuations in temperature and salinity, and a higher potential predation risk by the zooneuston. Furthermore, wind action promoting sea spray formation and bubbles rising from deeper water and bursting at the surface release SML-associated microbes into the atmosphere. In addition to being more concentrated compared to planktonic counterparts, the bacterioneuston, algae, and protists display distinctive community compositions compared to the underlying water, in both marine and freshwater habitats. Furthermore, the bacterial community composition was often dependent on the SML sampling device being used. While being well defined with respect to bacterial community composition, little is known about viruses in the SML, i.e., the virioneuston. This review has its focus on virus–bacterium dynamics at air–water interfaces, even if viruses likely interact with other SML microbes, including archaea and the phytoneuston, as can be deduced from viral interference with their planktonic counterparts. Although viruses were briefly mentioned as pivotal SML components in a recent review on this unique habitat, a synopsis of the emerging knowledge and the major research gaps regarding bacteriophages at air–water interfaces is still missing in the literature.