Impact of climate change and oligotrophication on quality and quantity of lake primary production: A case study in Lake Biwa.

Global climate change and anthropogenic oligotrophication are expected to reshape the dynamics of primary production (PP) in aquatic ecosystems; however, few studies have explored their long-term effects. In theory, the PP of phytoplankton in Lake Biwa may decline over decades due to warming, heightened stratification, and anthropogenic oligotrophication. Furthermore, the PP of large phytoplankton, which are inedible to zooplankton, along with biomass-specific productivity (PBc), could decrease. In this study, data from 1976 to 2021 and active fluorometry measurements taken in 2020 and 2021 were evaluated. Quantitatively, the temporal dynamics of mean seasonal PP during 1971-2021 were assessed according to the carbon fixation rate to investigate relationships among environmental factors. Qualitatively, phytoplankton biomass, PP, and PBc were measured in two size fractions [edible (S) or inedible (L) for zooplankton] in 2020 and 2021, and the L:S balance for these three measures was compared between 1992 (low-temperature/high-nutrient conditions) and 2020-2021 (high-temperature/low-nutrient conditions) to assess seasonal dynamics. The results indicated that climate change and anthropogenic oligotrophication over the past 30 years have diminished Lake Biwa's PP since the 1990s, impacting the phenology of PP dynamics. However, the L:S balance in PP and PBc has exhibited minimal change between the data from 1992 and the 2020-2021 period. These findings suggest that, although climate change and oligotrophication may reduce overall PP, they may not markedly alter the inedible/edible phytoplankton balance in terms of PP and PBc. Instead, as total PP declines, the production of small edible phytoplankton may decrease proportionally, potentially affecting trophic transfer efficiency and material cycling in Lake Biwa.

Measuring Photophysiology of Attached Stage of Colacium sp. by a Cuvette-Type Fast Repetition Rate Fluorometer.

Fast repetition rate fluorometer (FRRf) is a beneficial method for measuring photosystem II (PSII) photophysiology and primary productivity. Although FRRf can measure PSII absorption cross-section (σPSII), maximum photochemical efficiency (Fv/Fm), effective photochemical efficiency (Fq'/Fm'), and non-photochemical quenching (NPQNSV) for various eukaryotic algae and cyanobacteria, almost all FRRf studies to date have focused on phytoplankton. Here, the protocol describes how to measure PSII photophysiology of an epizoic alga Colacium sp. Ehrenberg 1834 (Euglenophyta), in its attached stage (attached to zooplankton), using cuvette-type FRRf. First, we estimated the effects of substrate zooplankton (Scapholeberis mucronata O.F. Müller 1776, Cladocera, Daphniidae) on baseline fluorescence and σPSII, Fv/Fm, Fq'/Fm', and NPQNSV of planktonic Colacium sp. To validate this methodology, we recorded photophysiology measurements of attached Colacium sp. on S. mucronata and compared these results with its planktonic stage. Representative results showed how the protocol could determine the effects of calcium (Ca) and manganese (Mn) on Colacium sp. photophysiology and identify the various effects of Mn enrichment between attached and planktonic stages. Finally, we discuss the adaptability of this protocol to other periphytic algae.

Development of photosynthetic carbon fixation model using multi-excitation wavelength fast repetition rate fluorometry in Lake Biwa.

Direct measurements of gross primary productivity (GPP) in the water column are essential, but can be spatially and temporally restrictive. Fast repetition rate fluorometry (FRRf) is a bio-optical technique based on chlorophyll a (Chl-a) fluorescence that can estimate the electron transport rate (ETRPSII) at photosystem II (PSII) of phytoplankton in real time. However, the derivation of phytoplankton GPP in carbon units from ETRPSII remains challenging because the electron requirement for carbon fixation (Фe,C), which is mechanistically 4 mol e- mol C-1 or above, can vary depending on multiple factors. In addition, FRRf studies are limited in freshwater lakes where phosphorus limitation and cyanobacterial blooms are common. The goal of the present study is to construct a robust Фe,C model for freshwater ecosystems using simultaneous measurements of ETRPSII by FRRf with multi-excitation wavelengths coupled with a traditional carbon fixation rate by the 13C method. The study was conducted in oligotrophic and mesotrophic parts of Lake Biwa from July 2018 to May 2019. The combination of excitation light at 444, 512 and 633 nm correctly estimated ETRPSII of cyanobacteria. The apparent range of Фe,C in the phytoplankton community was 1.1-31.0 mol e- mol C-1 during the study period. A generalised linear model showed that the best fit including 12 physicochemical and biological factors explained 67% of the variance in Фe,C. Among all factors, water temperature was the most significant, while photosynthetically active radiation intensity was not. This study quantifies the in situ FRRf method in a freshwater ecosystem, discusses core issues in the methodology to calculate Фe,C, and assesses the applicability of the method for lake GPP prediction.

A unified framework for herbivore-to-producer biomass ratio reveals the relative influence of four ecological factors.

The biomass ratio of herbivores to primary producers reflects the structure of a community. Four primary factors have been proposed to affect this ratio, including production rate, defense traits and nutrient contents of producers, and predation by carnivores. However, identifying the joint effects of these factors across natural communities has been elusive, in part because of the lack of a framework for examining their effects simultaneously. Here, we develop a framework based on Lotka-Volterra equations for examining the effects of these factors on the biomass ratio. We then utilize it to test if these factors simultaneously affect the biomass ratio of freshwater plankton communities. We found that all four factors contributed significantly to the biomass ratio, with carnivore abundance having the greatest effect, followed by producer stoichiometric nutrient content. Thus, the present framework should be useful for examining the multiple factors shaping various types of communities, both aquatic and terrestrial.

A shady phytoplankton paradox: when phytoplankton increases under low light.

Light is a fundamental driver of ecosystem dynamics, affecting the rate of photosynthesis and primary production. In spite of its importance, less is known about its community-scale effects on aquatic ecosystems compared with those of nutrient loading. Understanding light limitation is also important for ecosystem management, as human activities have been rapidly altering light availability to aquatic ecosystems. Here we show that decreasing light can paradoxically increase phytoplankton abundance in shallow lakes. Our results, based on field manipulation experiments, field observations and models, suggest that, under competition for light and nutrients between phytoplankton and submersed macrophytes, alternative stable states are possible under high-light supply. In a macrophyte-dominated state, as light decreases phytoplankton density increases, because macrophytes (which effectively compete for nutrients released from the sediment) are more severely affected by light reduction. Our results demonstrate how species interactions with spatial heterogeneity can cause an unexpected outcome in complex ecosystems. An implication of our findings is that partial surface shading for controlling harmful algal bloom may, counterintuitively, increase phytoplankton abundance by decreasing macrophytes. Therefore, to predict how shallow lake ecosystems respond to environmental perturbations, it is essential to consider effects of light on the interactions between pelagic and benthic producers.