The importance of plankton watching
The Sir Alister Hardy Foundation for Ocean Science (SAHFOS) in Plymouth UK has been collecting data on plankton from the North Atlantic and the North Sea since 1931 . The survey is named after an instrument, the Continuous Plankton Recorder (CPR), invented by Sir Alister Hardy, a brilliant marine biologist, perhaps better known to the public for his theory that humans evolved in the sea. He may still be right about human evolution, but it is his work on zooplankton and his foresight in establishing the CPR survey that should receive the highest accolade. The CRP survey is the longest running, large-scale marine biological survey in the world.
The CRP is towed by merchant ships on a number of designated shipping routes at a depth of about 10 m, collecting plankton on silk filters en route that are counted and analysed in the laboratory afterwards to generate a database, which now contains details on 400 species of plankton found in over 170 000 samples taken since 1946. This database has provided excellent material for researchers studying climate change and the impact of climate change on marine organisms.
SAHFOS researchers Anthony Richardson and Martin Edwards, and marine ecologist David Shoeman at the University of Port Elizabeth in South Africa, are among the scientists analysing this data most recently who have discovered abrupt changes in the plankton populations in the northeast Atlantic within the past decades that may have radically disrupted the food web, including commercial fisheries.
Phytoplankton primary production affected by climate change
Richardson and Shoeman  looked at more than a hundred taxonomic groups including phytoplankton diatoms and dinoflagellates, herbivorous zooplankton copepods that feed on phytoplankton, and carnivorous plankton such as arrow worms and amphipods that feed on the herbivores.
They found that changes in the abundance of phytoplankton correlated strongly with sea surface temperature, tending to be more abundant when cooler regions in the North Sea warmed, possibly because warming cold waters encouraged metabolism and growth, but decreased in abundance when warm regions in the North East Atlantic became even warmer, possibly because warm water blocks the upwelling of nutrient-rich deep water (Oceans and global warming, this series).
Richardson and Shoeman also found that while neither herbivore- nor carnivore-zooplankton abundance correlated with temperature, close correlations existed between the abundance of phytoplankton and herbivorous zooplankton, and between the abundance of herbivorous zooplankplanton and carnivorous zooplankton. This was evidence that the plankton food web is controlled “bottom-up” by the primary producers, and not “top-down” by the predators; and the effects of climate change on the primary producers could propagate up to fisheries.
The bottom-up control of the marine food web differs fundamentally from the terrestrial food web, and depends on the ability of phytoplankton to sequester carbon over the shortest timescales of any primary producers on earth. This gives phytoplankton a unique position not only in the marine food web, but also in the entire biosphere and the carbon cycle.
In terrestrial ecosystems, soil algae and cyanobacteria are important for maintaining soil fertility, controlling pollution and improving soil structure for plant growth , but are not directly at the bottom of the food web.
In the oceans, phytoplankton effectively funnels carbon into the biosphere to create an enormous standing carbon stock, and species that synthesize calcium carbonate shells also transport large amounts of carbon dioxide as they die and sink to the seabed where the shells remain as sediment for thousands of years. Fossil fuels are believed to have originated from plankton sediment that has been buried deep underground over millions of years (Oceans and global warming, this series). Thus, phytoplankton is the most important active carbon sink on earth, and without them, life in its current form is impossible.
Mismatch between plankton seasonal peaks
Transfer of carbon into the living biomass in the oceans depends on phytoplankton being consumed by larger organisms with longer life spans, which are in turn consumed by still larger organisms with even longer life spans and so on up to fish, seabirds, seals and whales. Efficient upward transfer of biomass depends on coupling between the peak seasons for consumer and the consumed, but that too appears to have become disturbed by global warming.
Richardson and Edwards  followed the shifts in the timing of seasonal plankton blooms (population peaks) over the past decades. Each species has an annual cycle of maximum and minimum, and zooplankton herbivores (feeding on the phytoplankton) and carnivores (feeding in turn on the herbivores) have evolved to exploit the phytoplankton bloom so as to transfer biomass effectively up the food web.
Since 1987, however, the cycles of different species have shifted out of synch, leading to a mismatch between the different tiers of plankton food web all considered ‘producers’ of the oceans. The phytoplankton diatoms and dinoflagellates are primary producers, the copepods, herbivores that feed on phytoplankton are secondary producers, non-copepod holozooplankton include both secondary producers and tertiary producers that feed on secondary producers, and meroplankton consist of fish larvae that are both secondary and tertiary producers.
The researchers found substantial shifts in the seasonal population peaks over the past few decades. In particular, the meroplankton peaks have shifted forward, echinoderms most of all by 47 days. In contrast, diatom peaks in spring and autumn have collectively remained unchanged, albeit with considerable variation among the species. Non-copepod holozooplankton show a wide range of responses, with some classes also occurring substantially earlier. The majority of dinoflagellates among the phytoplankton also occur earlier by 24 to 27 days.
For groups that peak when the water column is mixed or in a transitional state, there is considerable variability; whereas groups associated with low turbulent conditions have virtually all advanced in their seasonal peaks. The bottom-dwelling larvae of the meroplankton also showed larger shifts forward than their zooplankton prey.
During summer, meroplankton has moved forward collectively by 27 d, dinoflagellates by 23 days, copepods and non-copepod holozooplankton both by 10 days over the 45-year period. Diatoms as a group showed the largest variations with particular groups occurring both earlier and later during the spring and autumn peaks
In warm waters therefore, phytoplankton bloom occurred three weeks earlier, but the herbivores peaked only 10 days earlier. Thus, there could be less herbivores to feed the next level of predators, the fish larvae and carnivorous zooplankton.
“These effects at the base of the food web are so dramatic that they’re bound to have an effect on the whole North Atlantic ecology,” Edwards said .
Phytoplankton becomes unstable as surface waters warm
That’s not all. Jef Huisman of the University of Amsterdam, The Netherlands, and colleagues used a computer model to find out what happens to the phytoplankton when the surface waters of the oceans get warmer, and large parts of the oceans become stratified, as climate models predict; in other words, there is less vertical mixing as the lighter warm water sits on top of the dense cold water beneath.
Huisman and colleagues found that reduced vertical mixing caused major disturbances to phytoplankton growth, undermining the efficient sequestering of carbon dioxide and export of carbon to the deeper layers and the ocean floor. This could result in catastrophic positive feedback on global warming as carbon dioxide in the atmosphere increases, causing further increase to the sea surface temperature, more stratification of the water column, and further disturbance to phytoplankton growth.
Phytoplankton grows best at a depth of about 100 m, forming deep chlorophyll maxima (DCMs) in the oceans that reflect a compromise between light supplied from above and nutrients supplied from below. Scientists assumed these DCMs are stable features of the oceans.
However, Huisman and colleagues showed that reduced vertical mixing generate oscillations and chaos in phytoplankton biomass that destroy the DCMs. The two critical processes involved in the fluctuations are the sinking of dead plankton that removes nutrients from the surface layers where there is sufficient light for growth and the upward flux of nutrients required for new phytoplankton growth. The DCMs remain stable so long as the two processes are in balance. When vertical mixing is reduced beyond a critical threshold, nutrients in the top layers are depleted faster than they can be replaced, oscillations and chaos set in and the DCMs break up.
The model reproduces many real features. It predicts that DCMs form at a depth of about 100m, and span a similar range as in clear ocean waters (about 50 m). As consistent with observations, the model predicts that nutrients are depleted to near-zero levels above the DCM while the nutrient concentration increases linearly with depth below the DCM. Detailed time-series measurements from the subtropical North Pacific confirm the prediction of a vertical zoning of species, with different collections of species dominating at different depths, and seasonal patterns in chlorophyll and nutrient concentrations in the DCMs. The time series measurements also show additional fluctuations superimposed on the seasonal cycles, as predicted by the model. Also as predicted by the model, the time series measurements show that species with relatively high sinking velocities show larger variability than species with low sinking velocities.
These results are devastating enough, but they also converge with other studies which show that warming itself will directly undermine phytoplankton growth, quite apart from the stratification of oceans and the reduced supply of nutrients (Shutting down the oceans part III, global warming and plankton; snuffing out the green fuse, this series)
The Institute of Science in Society