Computational mechanisms for action selection

Imagine a zebra in the African savannah. At each moment in time this zebra has to weigh up alternative courses of action before deciding which will be most beneficial to it. For instance, it may want to graze because it is short of food, or it may want to head towards a water hole because it is short of water, or it may want to remain motionless in order to avoid detection by the predator it can see lurking nearby. This is an example of the problem of action selection: how to choose, at each moment in time, the most appropriate out of a repertoire of possible actions. This thesis investigates action selection in a novel way and makes three main contribu¬ tions. Firstly, a description is given of a simulated environment which is an extensive and detailed simulation of the problem of action selection for animals. Secondly, this simulated environment is used to investigate the adequacy of several theories of ac¬ tion selection such as the drive model, Lorenz's hydraulic model and Maes' spreading activation network. Thirdly, a new approach to action selection is developed which determines the most appropriate action in a principled way, and which does not suffer from the inherent shortcomings found in other methods

Defining the Action Selection Problem

There has been a lack of progress in the field of action selection due to an incomplete understanding of the problem being faced. The differing nature of constituent parts of the action selection/'timeallocation* problem has not been properly appreciated. Some common sub-problems, such as obtaining food and avoiding predators, are described in terms of the demands they make on an animal's time. The significant differences between these sub>- problems are highlighted and a classificatory scheme is proposed, with which sub-problems can be categorized. The need to take into account the full range of different sul)-problems is demonstrated with a few examples. A particular shortcoming shared by all of the more well-known action selection mechanisms, from both robotics and animal behaviour, is described

Geochemical evidence of denitrification in the Benguela upwelling system

This paper presents analysis of nitrate, phosphate and silicate data from the Benguela upwelling system. Evidence is presented that suggests denitrification occurring close to shore, and also nutrient trapping. Denitrification leaves an imprint on the water properties in terms of a nitrate deficit, that is to say nitrate concentrations that are significantly less than predicted by multiplying the phosphate concentrations by the Redfield ratio. It is probable that denitrification also causes a decoupling of nitrate and carbon compared to Redfield processes, and large-scale losses of nitrate in the Benguela which are not accompanied by losses of carbon. Nitrate-driven CO2 drawdown following upwelling will be less than it might otherwise be, because of denitrification. Nutrient trapping (secondary remineralisation) is apparent as enhanced phosphate concentrations, some of which are several Full-size image (<1 K) higher than in the offshore source waters for upwelling. Waters also become enriched in silicate and to a lesser extent nitrate as they advect across the shelf. By implication the same process should also “supercharge” waters in dissolved inorganic carbon, leading to stronger outgassing of CO2 immediately after upwelling. The effect is again to increase the size of the estimated Benguela upwelling system CO2 source

Analysis of longitudinal variations in North Pacific alkalinity to improve predictive algorithms

The causes of natural variation in alkalinity in the North Pacific surface ocean need to be investigated to understand the carbon cycle and to improve predictive algorithms. We used GLODAPv2 to test hypotheses on the causes of three longitudinal phenomena in Alk*, a tracer of calcium carbonate cycling. These phenomena are (a) an increase from east to west between 45°N and 55°N, (b) an increase from west to east between 25°N and 40°N, and (c) a minor increase from west to east in the equatorial upwelling region. Between 45°N and 55°N, Alk* is higher on the western than on the eastern side, and this is associated with denser isopycnals with higher Alk* lying at shallower depths. Between 25°N and 40°N, upwelling along the North American continental shelf causes higher Alk* in the east. Along the equator, a strong east-west trend was not observed, even though the upwelling on the eastern side of the basin is more intense, because the water brought to the surface is not high in Alk*. We created two algorithms to predict alkalinity, one for the entire Pacific Ocean north of 30°S and one for the eastern margin. The Pacific Ocean algorithm is more accurate than the commonly used algorithm published by Lee et al. (2006), of similar accuracy to the best previously published algorithm by Sasse et al. (2013), and is less biased with longitude than other algorithms in the subpolar North Pacific. Our eastern margin algorithm is more accurate than previously published algorithms

What drives the latitudinal gradient in open-ocean surface dissolved inorganic carbon concentration?

Previous work has not led to a clear understanding of the causes of spatial pattern in global surface ocean dissolved inorganic carbon (DIC), which generally increases polewards. Here, we revisit this question by investigating the drivers of observed latitudinal gradients in surface salinity-normalized DIC (nDIC) using the Global Ocean Data Analysis Project version 2 (GLODAPv2) database. We used the database to test three different hypotheses for the driver producing the observed increase in surface nDIC from low to high latitudes. These are (1) sea surface temperature, through its effect on the CO2 system equilibrium constants, (2) salinity-related total alkalinity (TA), and (3) highlatitude upwelling of DIC- and TA-rich deep waters. We find that temperature and upwelling are the two major drivers. TA effects generally oppose the observed gradient, except where higher values are introduced in upwelled waters. Temperature-driven effects explain the majority of the surface nDIC latitudinal gradient (182 of the 223 μmol kg1 increase from the tropics to the high-latitude Southern Ocean). Upwelling, which has not previously been considered as a major driver, additionally drives a substantial latitudinal gradient. Its immediate impact, prior to any induced air-sea CO2 exchange, is to raise Southern Ocean nDIC by 220 μmol kg1 above the average low-latitude value. However, this immediate effect is transitory. The long-term impact of upwelling (brought about by increasing TA), which would persist even if gas exchange were to return the surface ocean to the same CO2 as without upwelling, is to increase nDIC by 74 μmol kg1 above the low-latitude average

Phenological characteristics of global coccolithophore blooms

Coccolithophores are recognized as having a significant influence on the global carbon cycle through the production and export of calcium carbonate (often referred to as particulate inorganic carbon or PIC). Using remotely sensed PIC and chlorophyll data, we investigate the seasonal dynamics of coccolithophores relative to a mixed phytoplankton community. Seasonal variability in PIC, here considered to indicate changes in coccolithophore biomass, is identified across much of the global ocean. Blooms, which typically start in February–March in the low-latitude (~30°) Northern Hemisphere and last for ~6–7 months, get progressively later (April–May) and shorter (3–4 months) moving poleward. A similar pattern is observed in the Southern Hemisphere, where blooms that generally begin around August–September in the lower latitudes and which last for ~8 months get later and shorter with increasing latitude. It has previously been considered that phytoplankton blooms consist of a sequential succession of blooms of individual phytoplankton types. Comparison of PIC and chlorophyll peak dates suggests instead that in many open ocean regions, blooms of coccolithophores and other phytoplankton can co-occur, conflicting with the traditional view of species succession that is thought to take place in temperate regions such as the North Atlantic

Spatial variations in silicate-to-nitrate ratios in Southern Ocean surface waters are controlled in the short term by physics rather than biology

The nutrient composition (high in nitrate but low in silicate) of Subantarctic Mode Water (SAMW) forces diatom scarcity across much of the global surface ocean. This is because diatoms cannot grow without silicate. After formation and downwelling at the Southern Ocean's northern edge, SAMW re-emerges into the surface layers of the mid- and low-latitude oceans, providing a major nutrient source to primary producers in those regions. The distinctive nutrient composition of SAMW originates in the surface waters of the Southern Ocean, from which SAMW is formed. These waters are observed to transition from being rich in both silicate and nitrate in high-latitude areas of the Southern Ocean to being nitrate-rich but silicate-depleted at SAMW formation sites further north. Here we investigate the key controls of this change in nutrient composition with an idealised model, consisting of a chain of boxes linked by a residual (Ekman- and eddy-induced) overturning circulation. Biological processes are modelled on the basis of seasonal plankton bloom dynamics, and physical processes are modelled using a synthesis of outputs from the data-assimilative Southern Ocean State Estimate. Thus, as surface water flows northward across the Southern Ocean toward sites of SAMW formation, it is exposed in the model (as in reality) to seasonal cycles of both biology and physics. Our results challenge previous characterisations of the abrupt northward reduction in silicate-to-nitrate ratios in Southern Ocean surface waters as being predominantly driven by biological processes. Instead, our model indicates that, over shorter timescales (years to decades), physical processes connecting the deep and surface waters of the Southern Ocean (i.e. upwelling and entrainment) exert the primary control on the spatial distribution of surface nutrient ratios

Intercomparison of carbonate chemistry measurements on a cruise in northwestern European shelf seas

Four carbonate system variables were measured in surface waters during a cruise aimed at investigating ocean acidification impacts traversing northwestern European shelf seas in the summer of 2011. High-resolution surface water data were collected for partial pressure of carbon dioxide (pCO2; using two independent instruments) and pH using the total pH scale (pHT), in addition to discrete measurements of total alkalinity and dissolved inorganic carbon. We thus overdetermined the carbonate system (four measured variables, two degrees of freedom), which allowed us to evaluate the level of agreement between the variables on a cruise whose main aim was not intercomparison, and thus where conditions were more representative of normal working conditions. Calculations of carbonate system variables from other measurements generally compared well with direct observations of the same variables (Pearson’s correlation coefficient always greater than or equal to 0.94; mean residuals were similar to the respective accuracies of the measurements). We therefore conclude that four of the independent data sets of carbonate chemistry variables were of high quality. A diurnal cycle with a maximum amplitude of 41 μatm was observed in the difference between the pCO2 values obtained by the two independent analytical pCO2 systems, and this was partly attributed to irregular seawater flows to the equilibrator and partly to biological activity inside the seawater supply and one of the equilibrators. We discuss how these issues can be addressed to improve carbonate chemistry data quality on future research cruises

Environmental drivers of coccolithophore abundance and calcification across Drake Passage (Southern Ocean)

Although coccolithophores are not as numerically common or as diverse in the Southern Ocean as they are in subpolar waters of the North Atlantic, a few species, such as Emiliania huxleyi, are found during the summer months. Little is actually known about the calcite production (CP) of these communities or how their distribution and physiology relate to environmental variables in this region. In February 2009, we made observations across Drake Passage (between South America and the Antarctic Peninsula) of coccolithophore distribution, CP, primary production, chlorophyll a and macronutrient concentrations, irradiance and carbonate chemistry. Although CP represented less than 1% of total carbon fixation, coccolithophores were widespread across Drake Passage. The B/C morphotype of E. huxleyi was the dominant coccolithophore, with low estimates of coccolith calcite ( 0.01 pmol C coccolith-/ from biometric measurements. Both cell-normalised calcification (0.01–0.16 pmol C cell-1 d-1/ and total CP (< 20 μmol C m-1 d-1/were much lower than those observed in the subpolar North Atlantic where E. huxleyi morphotype A is dominant. However, estimates of coccolith production rates were similar (0.1–1.2 coccoliths cell-1 h-1/ to previous measurements made in the subpolar North Atlantic. A multivariate statistical approach found that temperature and irradiance together were best able to explain the observed variation in species distribution and abundance (Spearman’s rank correlation D0.4, p < 0.01). Rates of calcification per cell and coccolith production, as well as community CP and E. huxleyi abundance, were all positively correlated (p < 0.05) to the strong latitudinal gradient in temperature, irradiance and calcite saturation states across Drake Passage. Broadly, our results lend support to recent suggestions that coccolithophores, especially E. huxleyi, are advancing polewards. However, our in situ observations indicate that this may owe more to sea-surface warming and increasing irradiance rather than increasing CO2 concentrations

Geographical CO2 sensitivity of phytoplankton correlates with ocean buffer capacity

Accumulation of anthropogenic CO2 is significantly altering ocean chemistry. A range of biological impacts resulting from this oceanic CO2 accumulation are emerging, however the mechanisms responsible for observed differential susceptibility between organisms and across environmental settings remain obscure. A primary consequence of increased oceanic CO2 uptake is a decrease in the carbonate system buffer capacity, which characterises the system's chemical resilience to changes in CO2, generating the potential for enhanced variability in pCO2 and the concentration of carbonate [CO32‐], bicarbonate [HCO3‐] and protons [H+] in the future ocean. We conducted a meta‐analysis of 17 shipboard manipulation experiments performed across three distinct geographical regions that encompassed a wide range of environmental conditions from European temperate seas to Arctic and Southern oceans. These data demonstrated a correlation between the magnitude of natural phytoplankton community biological responses to short‐term CO2 changes and variability in the local buffer capacity across ocean basin scales. Specifically, short‐term suppression of small phytoplankton (<10 μm) net growth rates were consistently observed under enhanced pCO2 within experiments performed in regions with higher ambient buffer capacity. The results further highlight the relevance of phytoplankton cell size for the impacts of enhanced pCO2 in both the modern and future ocean. Specifically, cell‐size related acclimation and adaptation to regional environmental variability, as characterised by buffer capacity, likely influences interactions between primary producers and carbonate chemistry over a range of spatio‐temporal scales