Land use change and coastal water darkening drive synchronous dynamics in phytoplankton and fish phenology on centennial timescales.

At high latitudes, the suitable window for timing reproductive events is particularly narrow, promoting tight synchrony between trophic levels. Climate change may disrupt this synchrony due to diverging responses to temperature between, for example, the early life stages of higher trophic levels and their food resources. Evidence for this is equivocal, and the role of compensatory mechanisms is poorly understood. Here, we show how a combination of ocean warming and coastal water darkening drive long-term changes in phytoplankton spring bloom timing in Lofoten Norway, and how spawning time of Northeast Arctic cod responds in synchrony. Spring bloom timing was derived from hydrographical observations dating back to 1936, while cod spawning time was estimated from weekly fisheries catch and roe landing data since 1877. Our results suggest that land use change and freshwater run-off causing coastal water darkening has gradually delayed the spring bloom up to the late 1980s after which ocean warming has caused it to advance. The cod appear to track phytoplankton dynamics by timing gonadal development and spawning to maximize overlap between offspring hatch date and predicted resource availability. This finding emphasises the importance of land-ocean coupling for coastal ecosystem functioning, and the potential for fish to adapt through phenotypic plasticity.

Actions on sustainable food production and consumption for the post-2020 global biodiversity framework.

Current food production and consumption trends are inconsistent with the Convention on Biological Diversity's 2050 vision of living in harmony with nature. Here, we examine how, and under what conditions, the post-2020 biodiversity framework can support transformative change in food systems. Our analysis of actions proposed in four science-policy fora reveals that subsidy reform, valuation, food waste reduction, sustainability standards, life cycle assessments, sustainable diets, mainstreaming biodiversity, and strengthening governance can support more sustainable food production and consumption. By considering barriers and opportunities of implementing these actions in Peru and the United Kingdom, we derive potential targets and indicators for the post-2020 biodiversity framework. For targets to support transformation, genuine political commitment, accountability and compliance, and wider enabling conditions and actions by diverse agents are needed to shift food systems onto a sustainable path.

Centennial decline in North Sea water clarity causes strong delay in phytoplankton bloom timing.

With climate warming, a widespread expectation is that events in spring, such as flowering, bird migrations, and insect bursts, will occur earlier because of increasing temperature. At high latitudes, increased ocean temperature is suggested to advance the spring phytoplankton bloom due to earlier stabilization of the water column. However, climate warming is also expected to cause browning in lakes and rivers due to increases in terrestrial greening, ultimately reducing water clarity in coastal areas where freshwater drain. In shallow areas, decreased retention of sediments on the seabed will add to this effect. Both browning and resuspension of sediments imply a reduction of the euphotic zone and Sverdrup's critical depth leading to a delay in the spring bloom, counteracting the effect of increasing temperature. Here, we provide evidence that such a transparency reduction has already taken place in both the deep and shallow areas of the North Sea during the 20th century. A sensitivity analysis using a water column model suggests that the reduced transparency might have caused up to 3 weeks delay in the spring bloom over the last century. This delay stands in contrast to the earlier bloom onset expected from global warming, thus highlighting the importance of including changing water transparency in analyses of phytoplankton phenology and primary production. This appears to be of particular relevance for coastal waters, where increased concentrations of absorbing and scattering substances (sediments, dissolved organic matter) have been suggested to lead to coastal darkening.

Unclear associations between small pelagic fish and jellyfish in several major marine ecosystems.

During the last 20 years, a series of studies has suggested trends of increasing jellyfish (Cnidaria and Ctenophora) biomass in several major ecosystems worldwide. Some of these systems have been heavily fished, causing a decline among their historically dominant small pelagic fish stocks, or have experienced environmental shifts favouring jellyfish proliferation. Apparent reduction in fish abundance alongside increasing jellyfish abundance has led to hypotheses suggesting that jellyfish in these areas could be replacing small planktivorous fish through resource competition and/or through predation on early life stages of fish. In this study, we test these hypotheses using extended and published data of jellyfish, small pelagic fish and crustacean zooplankton biomass from four major ecosystems within the period of 1960 to 2014: the Southeastern Bering Sea, the Black Sea, the Northern California Current and the Northern Benguela. Except for a negative association between jellyfish and crustacean zooplankton in the Black Sea, we found no evidence of jellyfish biomass being related to the biomass of small pelagic fish nor to a common crustacean zooplankton resource. Calculations of the energy requirements of small pelagic fish and jellyfish stocks in the most recent years suggest that fish predation on crustacean zooplankton is 2-30 times higher than jellyfish predation, depending on ecosystem. However, compared with available historical data in the Southeastern Bering Sea and the Black Sea, it is evident that jellyfish have increased their share of the common resource, and that jellyfish can account for up to 30% of the combined fish-jellyfish energy consumption. We conclude that the best available time-series data do not suggest that jellyfish are outcompeting, or have replaced, small pelagic fish on a regional scale in any of the four investigated ecosystems. However, further clarification of the role of jellyfish requires higher-resolution spatial, temporal and taxonomic sampling of the pelagic community.

Light penetration structures the deep acoustic scattering layers in the global ocean.

The deep scattering layer (DSL) is a ubiquitous acoustic signature found across all oceans and arguably the dominant feature structuring the pelagic open ocean ecosystem. It is formed by mesopelagic fishes and pelagic invertebrates. The DSL animals are an important food source for marine megafauna and contribute to the biological carbon pump through the active flux of organic carbon transported in their daily vertical migrations. They occupy depths from 200 to 1000 m at daytime and migrate to a varying degree into surface waters at nighttime. Their daytime depth, which determines the migration amplitude, varies across the global ocean in concert with water mass properties, in particular the oxygen regime, but the causal underpinning of these correlations has been unclear. We present evidence that the broad variability in the oceanic DSL daytime depth observed during the Malaspina 2010 Circumnavigation Expedition is governed by variation in light penetration. We find that the DSL depth distribution conforms to a common optical depth layer across the global ocean and that a correlation between dissolved oxygen and light penetration provides a parsimonious explanation for the association of shallow DSL distributions with hypoxic waters. In enhancing understanding of this phenomenon, our results should improve the ability to predict and model the dynamics of one of the largest animal biomass components on earth, with key roles in the oceanic biological carbon pump and food web.

Jellyfish distribute vertically according to irradiance.

We tested the hypothesis that the coronate jellyfish Periphylla periphylla distributes vertically according to a preferential range of absolute light intensities. The study was carried out in Lurefjorden, Norway, a fjord characterized by mass occurrences of this jellyfish. We collected data on the vertical distribution of P. periphylla medusa during day, dusk and night periods from video observations by a remotely operated vehicle in relation to estimated ambient light levels. Our results suggest that large P. periphylla (average size in catches ~9 cm diameter) avoided total irradiance levels above 5×10-3 µmol quanta m-2 s-1. Nearly two-thirds of the population stayed above irradiance of 10-7 µmol quanta m-2 s-1 during daytime, while some individuals occupied much darker water. Thus, part of the population appeared to distribute vertically and undertake diel vertical migration (DVM) according to a preferential range of light intensities.

The emotion system promotes diversity and evolvability.

Studies on the relationship between the optimal phenotype and its environment have had limited focus on genotype-to-phenotype pathways and their evolutionary consequences. Here, we study how multi-layered trait architecture and its associated constraints prescribe diversity. Using an idealized model of the emotion system in fish, we find that trait architecture yields genetic and phenotypic diversity even in absence of frequency-dependent selection or environmental variation. That is, for a given environment, phenotype frequency distributions are predictable while gene pools are not. The conservation of phenotypic traits among these genetically different populations is due to the multi-layered trait architecture, in which one adaptation at a higher architectural level can be achieved by several different adaptations at a lower level. Our results emphasize the role of convergent evolution and the organismal level of selection. While trait architecture makes individuals more constrained than what has been assumed in optimization theory, the resulting populations are genetically more diverse and adaptable. The emotion system in animals may thus have evolved by natural selection because it simultaneously enhances three important functions, the behavioural robustness of individuals, the evolvability of gene pools and the rate of evolutionary innovation at several architectural levels.

Evidence of diel vertical migration in Mnemiopsis leidyi.

The vertical distribution and migration of plankton organisms may have a large impact on their horizontal dispersal and distribution, and consequently on trophic interactions. In this study we used video-net profiling to describe the fine scale vertical distribution of Mnemiopsis leidyi in the Kattegat and Baltic Proper. Potential diel vertical migration was also investigated by frequent filming during a 24-hour cycle at two contrasting locations with respect to salinity stratification. The video profiles revealed a pronounced diel vertical migration at one of the locations. However, only the small and medium size classes migrated, on average 0.85 m h(-1), corresponding to a total migration distance of 10 m during 12 h. Larger individuals (with well developed lobes, approx. >27 mm) stay on average in the same depth interval at all times. Biophysical data suggest that migrating individuals likely responded to light, and avoided irradiance levels higher than approx. 10 µmol quanta m(-2) s(-1). We suggest that strong stratification caused by low surface salinity seemed to prohibit vertical migration.

Effects of the emotion system on adaptive behavior.

A central simplifying assumption in evolutionary behavioral ecology has been that optimal behavior is unaffected by genetic or proximate constraints. Observations and experiments show otherwise, so that attention to decision architecture and mechanisms is needed. In psychology, the proximate constraints on decision making and the processes from perception to behavior are collectively described as the emotion system. We specify a model of the emotion system in fish that includes sensory input, neuronal computation, developmental modulation, and a global organismic state and restricts attention during decision making for behavioral outcomes. The model further includes food competition, safety in numbers, and a fluctuating environment. We find that emergent strategies in evolved populations include common emotional appraisal of sensory input related to fear and hunger and also include frequency-dependent rules for behavioral responses. Focused attention is at times more important than spatial behavior for growth and survival. Spatial segregation of the population is driven by personality differences. By coupling proximate and immediate influences on behavior with ultimate fitness consequences through the emotion system, this approach contributes to a unified perspective on the phenotype, by integrating effects of the environment, genetics, development, physiology, behavior, life history, and evolution.

Simulation of optically conditioned retention and mass occurrences of Periphylla periphylla.

Jellyfish blooms are of increasing concern in many parts of the world, and in Norwegian fjords an apparent increase in mass occurrences of the deep water jellyfish Periphylla periphylla has attracted attention. Here we investigate the hypothesis that changes in the water column light attenuation might cause local retention and thereby facilitate mass occurrences. We use a previously tested individual-based model of light-mediated vertical migration in P. periphylla to simulate how retention is affected by changes in light attenuation. Our results suggest that light attenuation, in combination with advection, has a two-sided effect on retention and that three fjord categories can be defined. In category 1, increased light attenuation turns fjords into dark "deep-sea" environments which increase the habitat and retention of P. periphylla. In category 2, an optimal light attenuation facilitates the maximum retention and likelihood for mass occurrences. In category 3, further increase in light attenuation, however, shoals the habitat so that individuals are increasingly exposed to advection and this results in loss of individuals and decreased retention. This classification requires accurate determinations of the organism's light preference, the water column light attenuation and topographical characteristics affecting advection.

Swimming and feeding in Periphylla periphylla (Scyphozoa, Coronatae).

Some western Norwegian fjords host extraordinarily abundant and persistent populations of the mesopelagic, coronate scyphomedusa, Periphylla periphylla. In these environments, from late autumn to spring, the medusae undertake regular diel vertical migrations into surface waters. From unique observations obtained with a remotely operated vehicle (ROV), including observations made without artificial light, we observed that 90% of the medusae swam with their tentacles in aboral position. Stomach content analyses of surface-collected specimens revealed that the medusae ate mainly calanoid copepods, but ostracods and large euphausiids were also prominent components of their diets. The clearance rate potential of P. periphylla, assessed from in situ observations and stomach contents, was comparable to that of similar-sized, epipelagic gelatinous species. Our findings suggest that P. periphylla behave as active predators in surface waters.

Endostyle cell recruitment as a frame of reference for development and growth in the Urochordate Oikopleura dioica.

In models of growth and life history, and in molecular and cell biology, there is a need for more accurate frames of reference to characterize developmental progression. In Caenorhabditis elegans, complete fate maps of cell lineage provide such a standard of reference. To be more widely applicable, reference frames should be easier to measure while still providing strong predictive capacity. Towards this aim, we have analyzed growth of the endostyle in the appendicularian Oikopleura dioica at the cellular level, and measured its response to temperature and food availability. Specifically, we test the hypothesis that age of a specific developmental stage in O. dioica can be predicted from the number of endostyle cells and temperature. We show that the endostyle grows by recruiting cells from the posterior tip into the lateral arms of the organ in an anterior-posterior orientation and that the rate of increase in lateral arm endostyle cells is temperature-dependent but unresponsive to nutritional intake. Endostyle cells therefore serve as an accurate and easily measured marker to describe developmental progression. Conceptually, such a method of characterizing developmental progression should help bridge life-history events and molecular mechanisms throughout organismal aging, facilitating cross-disciplinary understanding by providing a common experimental framework.