Shovonlal Roy

The stability of ecosystems: A brief overview of the paradox of enrichment

In theory, enrichment of resource in a predator-prey model leads to destabilization of the system, thereby collapsing the trophic interaction, a phenomenon referred to as “the paradox of enrichment”. After it was first proposed by Rosenzweig (1971), a number of subsequent studies were carried out on this dilemma over many decades. In this article, we review these theoretical and experimental works and give a brief overview of the proposed solutions to the paradox. The mechanisms that have been discussed are modifications of simple predator-prey models in the presence of prey that is inedible, invulnerable, unpalatable and toxic. Another class of mechanisms includes an incorporation of a ratio-dependent functional form, inducible defence of prey and density-dependent mortality of the predator. Moreover, we find a third set of explanations based on complex population dynamics including chaos in space and time. We conclude that, although any one of the various mechanisms proposed so far might potentially prevent destabilization of the predator-prey dynamics following enrichment, in nature different mechanisms may combine to cause stability, even when a system is enriched. The exact mechanisms, which may differ among systems, need to be disentangled through extensive field studies and laboratory experiments coupled with realistic theoretical models.

A Consumer's Guide to Satellite Remote Sensing of Multiple Phytoplankton Groups in the Global Ocean

Phytoplankton are composed of diverse taxonomical groups, which are manifested as distinct morphology, size and pigment composition. These characteristics, modulated by their physiological state, impact their light absorption and scattering, allowing them to be detected with ocean color satellite radiometry. There is a growing volume of literature describing satellite algorithms to retrieve information on phytoplankton composition in the ocean. This synthesis provides a review of current methods and a simplified comparison of approaches. The aim is to provide an easily comprehensible resource for non-algorithm developers, who desire to use these products, thereby raising the level of awareness and use of these products and reducing the boundary of expert knowledge needed to make a pragmatic selection of output products with confidence. The satellite input and output products, their associated validation metrics, as well as assumptions, strengths and limitations of the various algorithm types are described, providing a framework for algorithm organization to assist users and inspire new aspects of algorithm development capable of exploiting the higher spectral, spatial and temporal resolutions from the next generation of ocean color satellites.

Retrieval of phytoplankton size from bio-optical measurements: theory and applications

The absorption coefficient of a substance distributed as discrete particles in suspension is less than that of the same material dissolved uniformly in a medium—a phenomenon commonly referred to as the flattening effect. The decrease in the absorption coefficient owing to flattening effect depends on the concentration of the absorbing pigment inside the particle, the specific absorption coefficient of the pigment within the particle, and on the diameter of the particle, if the particles are assumed to be spherical. For phytoplankton cells in the ocean, with diameters ranging from less than 1 µm to more than 100 µm, the flattening effect is variable, and sometimes pronounced, as has been well documented in the literature. Here, we demonstrate how the in vivo absorption coefficient of phytoplankton cells per unit concentration of its major pigment, chlorophyll a, can be used to determine the average cell size of the phytoplankton population. Sensitivity analyses are carried out to evaluate the errors in the estimated diameter owing to potential errors in the model assumptions. Cell sizes computed for field samples using the model are compared qualitatively with indirect estimates of size classes derived from high performance liquid chromatography data. Also, the results are compared quantitatively against measurements of cell size in laboratory cultures. The method developed is easy-to-apply as an operational tool for in situ observations, and has the potential for application to remote sensing of ocean colour data.

Toxin-allelopathy among phytoplankton species prevents competitive exclusion

Toxic or allelopathic compounds liberated by toxin-producing phytoplankton (TPP) acts as a strong mediator in plankton dynamics. On an analysis of a set of phytoplankton biomass data that have been collected by our group in the northwest part of the Bay of Bengal, and by analysis of a three-component mathematical model under a constant as well as a stochastic environment, we explore the role of toxin-allelopathy in determining the dynamic behavior of the competing phytoplankton species. The overall results, based on analytical and numerical wings, demonstrate that toxin-allelopathy due to the TPP promotes a stable co-existence of those competitive phytoplankton that would otherwise exhibit competitive exclusion of the weak species. Our study suggests that TPP might be a potential candidate for maintaining the co-existence and diversity of competing phytoplankton species.

Enrichment and ecosystem stability: effect of toxic food.

Enrichment in resource availability theoretically destabilizes predator-prey dynamics (the paradox of enrichment). However, a minor change in the resource stoichiometry may make a prey toxic for the predator, and the presence of toxic prey affects the dynamics significantly. Here, theoretically we explore how, at increased carrying capacity, a toxic prey affects the oscillation or destabilization of predator-prey dynamics, and how its presence influences the growth of the predator as well as that of a palatable prey. Mathematical analysis determines the bounds on the food toxicity that allow the coexistence of a predator along with a palatable and a toxic prey. The overall results demonstrate that toxic food counteracts oscillation (destabilization) arising from enrichment of resource availability. Moreover, our results show that, at increased resource availability, toxic food that acts as a source of extra mortality may increase the abundance of the predator as well as that of the palatable prey.

Interaction among non-toxic phytoplankton, toxic phytoplankton and zooplankton: inferences from field observations.

We explore the mutual dependencies and interactions among different groups of species of the plankton population, based on an analysis of the long-term field observations carried out by our group in the North–West coast of the Bay of Bengal. The plankton community is structured into three groups of species, namely, non-toxic phytoplankton (NTP), toxic phytoplankton (TPP) and zooplankton. To find the pair-wise dependencies among the three groups of plankton, Pearson and partial correlation coefficients are calculated. To explore the simultaneous interaction among all the three groups, a time series analysis is performed. Following an Expectation Maximization (E-M) algorithm, those data points which are missing due to irregularities in sampling are estimated, and with the completed data set a Vector Auto-Regressive (VAR) model is analyzed. The overall analysis demonstrates that toxin-producing phytoplankton play two distinct roles: the inhibition on consumption of toxic substances reduces the abundance of zooplankton, and the toxic materials released by TPP significantly compensate for the competitive disadvantages among phytoplankton species. Our study suggests that the presence of TPP might be a possible cause for the generation of a complex interaction among the large number of phytoplankton and zooplankton species that might be responsible for the prolonged coexistence of the plankton species in a fluctuating biomass.

Effect of disease-selective predation on prey infected by contact and external sources

We propose and analyze a simple mathematical model for susceptible prey (S)-infected prey (I)-predator (P) interaction, where the susceptible prey population (S) is infected directly from external sources as well as through contact with infected class (I) and the predator completely avoids consuming the infected prey. The model is analyzed to obtain different thresholds of the key parameters under which the system exhibits stability around the biologically feasible equilibria. Through numerical simulations we display the effects of external infection and the infection through contact on the system dynamics in the absence as well as in the presence of the predator. We compare the system dynamics when infection occurs only through contact, with that when it occurs through contact and external sources. Our analysis demonstrates that under a disease-selective predation, stability and oscillations of the system is determined by two key parameters: the external infection rate and the force of infection through contact. Due to the introduction of external infection, the predator and the prey population show limit-cycle oscillations over a range parametric values. We suggest that while predicting the dynamics of such an eco-epidemiological system, the modes of infection and the infection rates might be carefully investigated.

Spatial interaction among nontoxic phytoplankton, toxic phytoplankton, and zooplankton: emergence in space and time.

In homogeneous environments, by overturning the possibility of competitive exclusion among phytoplankton species, and by regulating the dynamics of overall plankton population, toxin-producing phytoplankton (TPP) potentially help in maintaining plankton diversity—a result shown recently. Here, I explore the competitive effects of TPP on phytoplankton and zooplankton species undergoing spatial movements in the subsurface water. The spatial interactions among the species are represented in the form of reaction-diffusion equations. Suitable parametric conditions under which Turing patterns may or may not evolve are investigated. Spatiotemporal distributions of species biomass are simulated using the diffusivity assumptions realistic for natural planktonic systems. The study demonstrates that spatial movements of planktonic systems in the presence of TPP generate and maintain inhomogeneous biomass distribution of competing phytoplankton, as well as grazer zooplankton, thereby ensuring the persistence of multiple species in space and time. The overall results may potentially explain the sustainability of biodiversity and the spatiotemporal emergence of phytoplankton and zooplankton species under the influence of TPP combined with their physical movement in the subsurface water.

Modeling migratory grazing of zooplankton on toxic and non-toxic phytoplankton

Migratory grazing of zooplankton between non-toxic phytoplankton (NTP) and toxic phytoplankton (TPP) is a realistic phenomena unexplored so far. The present article is a first step in this direction. A mathematical model of NTP–TPP-zooplankton with constant and variable zooplankton migration is proposed and analyzed. The asymptotic dynamics of the model system around the biologically feasible equilibria is explored through local stability analysis. The dynamics of the proposed system is explored and displayed for different combination of migratory parameters and toxin inhibition parameters. Our analysis suggests that the migratory grazing of zooplankton has a significant role in determining the dynamic stability and oscillation of phytoplankton zooplankton systems.

Out of Amazonia: Late-Holocene climate change and the Tupi–Guarani trans-continental expansion

The late-Holocene expansion of the Tupi–Guarani languages from southern Amazonia to SE South America constitutes one of the largest expansions of any linguistic family in the world, spanning ~4000 km between latitudes 0°S and 35°S at about 2.5k cal. yr BP. However, the underlying reasons for this expansion are a matter of debate. Here, we compare continental-scale palaeoecological, palaeoclimate and archaeological datasets, to examine the role of climate change in facilitating the expansion of this forest-farming culture. Because this expansion lies within the path of the South American Low-Level Jet, the key mechanism for moisture transport across lowland South America, we were able to explore the relationship between climate change, forest expansion and the Tupi–Guarani. Our data synthesis shows broad synchrony between late-Holocene increasing precipitation and southerly expansion of both tropical forest and Guarani archaeological sites – the southernmost branch of the Tupi–Guarani. We conclude that climate change likely facilitated the agricultural expansion of the Guarani forest-farming culture by increasing the area of forested landscape that they could exploit, showing a prime example of ecological opportunism.