Bioluminescent Dinoflagellates Research Articles

Environmental controls on bioluminescent dinoflagellate density, in Laguna Grande, Fajardo, Puerto Rico

Introduction: Bio bays in Puerto Rico play an important socio-economic role and declines in dominant bioluminescent dinoflagellate Pyrodinium bahamense are concerning. Studies show erratic blooms with is weak correlation to in situ environmental factors. Our study examines shorter field and longer proxy records on dinoflagellate density at Laguna Grande de Fajardo (LGF). Objectives: To quantify temporal changes in dinoflagellate density in a long-term monitoring study, understand how the marine environment modulates those changes, and determine the wider impacts of a fluctuating climate and extreme events on proxies for dinoflagellate density. Methods: Bimonthly samples were collected from 2016 to 2021 at three sampling sites in LGF. Dinoflagellates density was estimated by Sedgewick Rafter counting cells. Environmental conditions were obtained from Rio Fajardo 5007100 station and NOAA buoy 41056. Marine climate and biotic proxies were obtained from remote sensing measurements. Kruskal Wallis, Spearman correlations and cross-correlations in the shorter field and longer proxy records were used to evaluate environmental controls on LGF dinoflagellate blooms. Results: Six years of field monitoring densities found a low period in 2016-2017, frequent and intense blooms in 2018-2021 punctuated by hurricanes. Generally low values were recorded in late winter in contrast with higher values in late summer (Aug-Nov). Light winds and mixed layer response to seasonal warming in the form of high tides and low salinity, were found to sustain dinoflagellate reproduction. Conclusions: Bioluminescent dinoflagellates are vital to coastal tourism and require resource management. LGF results show that: 1) dinoflagellate counts fluctuate widely, 2) fluorescing dinoflagellates are sensitive to environmental conditions because of limited seasonality and narrow physiological range, 3) hurricanes play a role by ‘raking and refreshing’ the coastal lagoon for subsequent biotic reproduction, and 4) intra-seasonal fluctuations of density and proxies relate to air-sea thermodynamic conditions, the salinity budget and sea level.

Toxin Dynamics among Populations of the Bioluminescent HAB Species Pyrodinium bahamense from the Indian River Lagoon, FL.

Dinoflagellate species that form some of the most frequent toxic blooms are also bioluminescent, yet the two traits are rarely linked when studying bloom development and persistence. P. bahamense is a toxic, bioluminescent dinoflagellate that previously bloomed in Florida with no known record of saxitoxin (STX) production. Over the past 20 years, STX was identified in P. bahamense populations. The goal of this study was to examine toxin dynamics and associated molecular mechanisms in spatially and temporally distinct P. bahamense populations from the Indian River Lagoon, FL. SxtA4 is a key gene required for toxin biosynthesis. SxtA4 genotype analysis was performed on individual cells from multiple sites. Cell abundance, toxin quota cell-1, and sxtA4 and RubisCo (rbcL) transcript abundance were also measured. There was a significant negative correlation between cell abundance and toxin quota cell-1. While the sxtA4+ genotype was dominant at all sites, its frequency varied, but it occurred at 90-100% in many samples. The underlying mechanism for toxin decrease with increased cell abundance remains unknown. However, a strong, statistically significant negative correlation was found between stxA4 transcripts and the sxtA4/rbcL ratio, suggesting cells make fewer sxtA4 transcripts as a bloom progresses. However, the influence of sxtA4- cells must also be considered. Future plans include bioluminescence measurements, normalized to a per cell basis, at sites when toxicity is measured along with concomitant quantification of sxtA4 gene and transcript copy numbers as a means to elucidate whether changes in bloom toxicity are driven more at the genetic (emergence of sxtA4- cells) or transcriptional (repression of sxtA4 in sxtA4+ cells) level. Based on the results of this study, a model is proposed that links the combined traits of toxicity and bioluminescence in P. bahamense bloom development.

Effect of salinity on the bioluminescence intensity of the heterotrophic dinoflagellates Noctiluca scintillans and Polykrikos kofoidii and the autotrophic dinoflagellate Alexandrium mediterraneum

Many dinoflagellate species are bioluminescent, which is one of the anti-predation mechanisms in these species. In addition, dinoflagellate species experience a wide range of salinities in the ocean. However, the effects of salinity on their bioluminescence intensity has only been investigated for one species. Here, we explored the effect of salinity on the bioluminescence intensity of the heterotrophic dinoflagellate Noctiluca scintillans NSDJ2010 feeding on the chlorophyte Dunaliella salina, the heterotrophic dinoflagellate Polykrikos kofoidii PKJH1607 feeding on the dinoflagellate Alexadrium minutum, and the autotrophic dinoflagellate Alexandrium mediterraneum AMYS1807. Moreover, to determine the cell volume and growth effects on bioluminescence intensity, the cell volume and growth rate of three bioluminescent dinoflagellates were simultaneously investigated. The mean 200-s-integrated bioluminescence intensity (BL) per cell, equivalent to the total bioluminescence, of N. scintillans, P. kofoidii, and A. mediterraneum was significantly affected by salinity and increased with increasing salinity from 10 to 40. The results of the present study suggest that the total bioluminescence of N. scintillans, P. kofoidii, and A. mediterraneum in offshore and oceanic waters is greater than that in estuarine waters.

Bioluminescence - The Vibrant Glow of Nature and its Chemical Mechanisms.

Bioluminescence, the mesmerizing natural phenomenon where living organisms produce light through chemical reactions, has long captivated scientists and laypersons alike, offering a rich tapestry of insights into biological function, ecology, evolution as well as the underlying chemistry. This comprehensive introductory review systematically explores the phenomenon of bioluminescence, addressing its historical context, geographic dispersion, and ecological significance with a focus on their chemical mechanisms. Our examination begins with terrestrial bioluminescence, discussing organisms from different habitats. We analyze thefireflies of Central Europe's meadows and the fungi in the Atlantic rainforest of Brazil. Additionally, we inspect bioluminescent species in New Zealand, specifically river-dwelling snails and mosquito larvae found in Waitomo Caves. Our exploration concludes in the Siberian Steppes, highlighting the area's luminescent insects and annelids. Transitioning to the marine realm, the second part of this review examines marine bioluminescent organisms. We explore this phenomenon in deep-sea jellyfish and their role in the ecosystem. We then move to Toyama Bay, Japan, where seasonal bioluminescence of dinoflagellates and ostracods present a unique case study. We also delve into the bacterial world, discussing how bioluminescent bacteria contribute to symbiotic relationships. For each organism, we contextualize its bioluminescence, providing details about its discovery, ecological function, and geographical distribution. A special focus lies on the examination of the underlying chemical mechanisms that enables these biological light displays. Concluding this review, we present a series of practical bioluminescence and chemiluminescence experiments, providing a resource for educational demonstrations and student research projects. Our goal with this review is to provide a summary of bioluminescence across the diverse ecological contexts, contributing to the broader understanding of this unique biological phenomenon and its chemical mechanisms serving researchers new to the field, educators and students alike.

New investigation of the cyst–motile relationship for Votadinium spinosum reveals a Protoperidinium claudicans species complex (Dinophyceae, Peridiniales)

Protoperidinium claudicans is a planktonic, heterotrophic, bioluminescent dinoflagellate species commonly found in neritic waters. It has long been considered to display phenotypic variation in its second anterior intercalary plate, which could vary between quadra, penta and hexa. The equivalent spinose, cordate cyst goes under the name of Votadinium spinosum. Here we perform incubation experiments with cysts from France, Canada, China and Japan, which demonstrate that P. claudicans forms a species complex, with at least two ribotypes with a penta configuration (P. claudicans) and one with a quadra configuration (P. carriae sp. nov.). A fossil-based cyst, V. multispinosum sp. nov., is described as the equivalent of P. carriae. Molecular phylogenetics using large subunit ribosomal DNA supports these observations. The cyst–theca relationships for Votadinium psilodora and another, undescribed, cordate, spineless Votadinium species are also reported from China. Macromolecular analyses of the cyst wall of V. multispinosum reveal it is comprised of a protein-rich carbohydrate compound. We show that this compound is not uncommon in dinoflagellate and ciliate cysts and that it is unlikely to preserve very well upon sedimentation and burial.

Perspectives on marine dinoflagellate bioluminescence: Is the current prevalence along the Indian waters only reckoned with aesthetic pleasure?

Perspectives on marine dinoflagellate bioluminescence: Is the current prevalence along the Indian waters only reckoned with aesthetic pleasure?

Revisiting the burglar alarm hypothesis: A behavioural cascade mediated by dinoflagellate bioluminescence

Abstract Bioluminescence is widespread among marine organisms and has evolved independently multiple times. While its specific adaptive value is diverse, bioluminescence in most cases mediates fundamental interactions between individuals (predator, prey, mates) and thus impacts ecosystem processes. One hypothesized value of bioluminescence in dinoflagellates is through the ‘burglar alarm’: Grazers of phytoplankton will make the ambient water ‘glow’ as they swim, thereby attracting visual predators of the grazer, thus indirectly protecting the dinoflagellates. However, the most important grazers of dinoflagellates, copepods, are generally too small to elicit dinoflagellates to glow. Only individual cells captured by a copepod will flash, which in turn elicits a powerful escape response in the copepod. Here, we test a variant of this hypothesis that may work for copepods. The behavioural response of the grazer to the flashing of a captured dinoflagellate, rather than the flashing itself, attracts the attention of the grazer's flow‐sensing predators. We demonstrate that bioluminescence in three dinoflagellates reduces the clearance and ingestion rate of nauplii of the copepod Temora longicornis. The presence of bioluminescent cells also elicited an increased frequency of high‐speed jumps of the grazers. The increased jump frequency elevated the detectability of the grazers to a flow‐sensing predator, the copepod Centropages typicus, consequently leading to increased predation mortality of T. longicornis nauplii. The consequent behavioural cascade mediated by bioluminescence works for small grazers that cause only single cells to flash, unlike the traditional description of the burglar alarm. The significance of such behavioural mediated indirect interactions in shaping food webs is becoming increasingly recognized, although rarely demonstrated in pelagic food webs. We argue that the burglar alarm is a secondary adaptation, while the defence of individual cells is the primary adaptive value of bioluminescence in dinoflagellates. Read the free Plain Language Summary for this article on the Journal blog.

Role of voltage-gated proton channel (Hv1) in cancer biology.

The acid-base characteristics of tumor cells and the other elements that compose the tumor microenvironment have been topics of scientific interest in oncological research. There is much evidence confirming that pH conditions are maintained by changes in the patterns of expression of certain proton transporters. In the past decade, the voltage-gated proton channel (Hv1) has been added to this list and is increasingly being recognized as a target with onco-therapeutic potential. The Hv1 channel is key to proton extrusion for maintaining a balanced cytosolic pH. This protein-channel is expressed in a myriad of tissues and cell lineages whose functions vary from producing bioluminescence in dinoflagellates to alkalizing spermatozoa cytoplasm for reproduction, and regulating the respiratory burst for immune system response. It is no wonder that in acidic environments such as the tumor microenvironment, an exacerbated expression and function of this channel has been reported. Indeed, multiple studies have revealed a strong relationship between pH balance, cancer development, and the overexpression of the Hv1 channel, being proposed as a marker for malignancy in cancer. In this review, we present data that supports the idea that the Hv1 channel plays a significant role in cancer by maintaining pH conditions that favor the development of malignancy features in solid tumor models. With the antecedents presented in this bibliographic report, we want to strengthen the idea that the Hv1 proton channel is an excellent therapeutic strategy to counter the development of solid tumors.

AWWA Water Science Author Spotlight

<i>AWWA Water Science</i> Author Spotlight

Bioluminescent Dinoflagellates as a Bioassay for Toxicity Assessment.

Dinoflagellates bioluminescence mechanism depends upon a luciferin-luciferase reaction that promotes blue light emission (480 nm) in specialized luminogenic organelles called scintillons. The scintillons contain luciferin, luciferase and, in some cases, a luciferin-binding protein (LBP), which prevents luciferin from non-enzymatic oxidation in vivo. Even though dinoflagellate bioluminescence has been studied since the 1950s, there is still a lack of mechanistic understanding on whether the light emission process involves a peroxidic intermediate or not. Still, bioassays employing luminous dinoflagellates, usually from Gonyaulax or Pyrocystis genus, can be used to assess the toxicity of metals or organic compounds. In these dinoflagellates, the response to toxicity is observed as a change in luminescence, which is linked to cellular respiration. As a result, these changes can be used to calculate a percentage of light inhibition that correlates directly with toxicity. This current approach, which lies in between fast bacterial assays and more complex toxicity tests involving vertebrates and invertebrates, can provide a valuable tool for detecting certain pollutants, e.g., metals, in marine sediment and seawater. Thus, the present review focuses on how the dinoflagellates bioluminescence can be applied to evaluate the risks caused by contaminants in the marine environment.

Thorough Understanding of Bioluminophore Production in Bacterial Bioluminescence.

Bioluminophore of bioluminescence (BL) comes from the decomposition of peroxide, which is an intermediate produced in the complicated chemical reactions of BL. The peroxide is a dioxetanone in most BL cases and an endoperoxide in fungal BL. The decomposition mechanisms of these two types of peroxides have been exclusively studied. However, the peroxide is a linear organic peroxide in bacterial BL, whose decomposition explanations are quite controversial, and seven mechanisms have been proposed. To thoroughly understand the mechanism of bioluminophore production in bacterial BL, this paper systematically discusses the seven proposed mechanisms via the present computational results and previous experimental and theoretical results. Our research results indicate that the bioluminophore in bacterial BL is produced through the charge-transfer initiated luminescence (CTIL) mechanism. The decomposition mechanism of linear organic peroxide was compared with the decomposition mechanisms of the other two types of peroxides, dioxetanone and endoperoxide. This study is also helpful in understanding the bioluminophore production in other BLs via the decomposition of an organic peroxide, such as dinoflagellate BL.

Factors responsible for the sudden outburst of Noctiluca scintillans in the Chennai coastal waters, southeast coast of India – a case study

The paper discusses the factors associated with the sudden outbreak of the nocturnal heterotrophic bioluminescent dinoflagellate Noctiluca scintillans along the Chennai coast. The bloom occurred along a stretch of 16 km following a spell of heavy rain in August 2019. The density of N. scintillans varied from 1000 to 19000 cells/L, with a distinguished distribution pattern. High N. scintillans abundance was recorded at Panaiyurkuppam and Kovalam, with 19000 cells/L and 18000 cells/L, recorded respectively. Adequate nutrients brought by substantial rainfall and a high abundance of the diatom Thalassiosira sp. triggered the N. scintillans bloom. The low wind speed (5 m/sec), lowering of atmospheric temperature (from 24.00 to 31.00°C, 27.5 ± 1.17°C), high rainfall (6 mm within one day), and low sea surface temperature (SST) (from 25.20 to 31.00°C, 29.37± 1.17°C) are probable environmental cues. Local hydrodynamics and the diverging currents governed the presence and dispersion of the bloom in the region.

Detection of bioluminescent dinoflagellates based on luciferase genes during the ‘blue tears’ season around the Matsu archipelago

Noctiluca scintillans is considered to be a bioluminescent bloom-forming species in the coastal water around the Matsu archipelago. To identify the bioluminescent dinoflagellates and their distributions around the Matsu archipelago, metatranscriptome and luciferase (lcf) gene sequencing were conducted from June 2016 to July 2017. Metatranscriptomes retrieved lcf genes mainly from Noctiluca and other bioluminescent dinoflagellates. This result demonstrates that lcf genes were actually expressed in multiple dinoflagellate species. An analysis of the lcf composition of dinoflagellates indicated that N. scintillans was the dominant bioluminescent species during May and July. In late summer, this dominance was replaced by other bioluminescent dinoflagellate species, such as Alexandrium affine and Ceratium fusus. No lcf gene from the known toxic bioluminescent dinoflagellates was obtained during the period of investigation. Our results suggest that N. scintillans is not the only dinoflagellate species producing bioluminescence around the Matsu archipelago.

Night on Earth

Night on Earth

Sexual Reproduction in Dinoflagellates—The Case of Noctiluca scintillans and Its Ecological Implications

Noctiluca scintillans is a larger, bioluminescent red-tide dinoflagellate (400–1,000 μm in diameter) that reproduces by sexual or asexual reproduction (binary fission). The process of sexual reproduction in N. scintillans has been thoroughly studied, but the ecological role and the mechanism of shifting from asexual to sexual reproduction have not been fully elucidated. It is believed, however, that sexual reproduction occurs when N. scintillans faces environmental stress. In this study, we tried to determine which factors drive N. scintillans to undergo sexual reproduction and we considered sexual reproduction’s ecological role. We cultured N. scintillans under different conditions of temperature, N. scintillans cell concentration, prey concentration, cultivation time, cultivation volume, light exposure time and physical vibration (simulated wave motion), and counted gametocyte mother cells every 24 h to calculate how the sexual reproduction rate changed over the experimental period. Rises in the sexual reproduction rate or the concentration of gametocyte mother cells only occurred in response to large variations in prey concentration, typically after the exponential phase of N. scintillans population growth. A noticeable upsurge in gametocyte mother cells, from 1% or less to nearly 10% of the total N. scintillans population, occurred when the prey concentration fell below ∼400 cells/mL. This implies that a sudden decrease in prey concentration induces more N. scintillans to shift from trophonts to gametocyte mother cells. We suggest that sexual reproduction may occur in N. scintillans as a response to the post-bloom situation when the dinoflagellate’s food supply has been dramatically depleted, producing large numbers of gametes for an alternative mode of survival after the end of each bloom.

SxtA4+ and sxtA4- Genotypes Occur Together within Natural Pyrodinium bahamense Sub-Populations from the Western Atlantic.

Saxitoxin (STX) is a secondary metabolite and potent neurotoxin produced by several genera of harmful algal bloom (HAB) marine dinoflagellates. The basis for variability in STX production within natural bloom populations is undefined as both toxic and non-toxic strains (of the same species) have been isolated from the same geographic locations. Pyrodinium bahamense is a STX-producing bioluminescent dinoflagellate that blooms along the east coast of Florida as well as the bioluminescent bays in Puerto Rico (PR), though no toxicity reports exist for PR populations. The core genes in the dinoflagellate STX biosynthetic pathway have been identified, and the sxtA4 gene is essential for toxin production. Using sxtA4 as a molecular proxy for the genetic capacity of STX production, we examined sxtA4+ and sxtA4- genotype frequency at the single cell level in P. bahamense populations from different locations in the Indian River Lagoon (IRL), FL, and Mosquito Bay (MB), a bioluminescent bay in PR. Multiplex PCR was performed on individual cells with Pyrodinium-specific primers targeting the 18S rRNA gene and sxtA4. The results reveal that within discrete natural populations of P. bahamense, both sxtA4+ and sxtA4- genotypes occur, and the sxtA4+ genotype dominates. In the IRL, the frequency of the sxtA4+ genotype ranged from ca. 80–100%. In MB, sxtA4+ genotype frequency ranged from ca 40–66%. To assess the extent of sxtA4 variation within individual cells, sxtA4 amplicons from single cells representative of the different sampling sites were cloned and sequenced. Overall, two variants were consistently obtained, one of which is likely a pseudogene based on alignment with cDNA sequences. These are the first data demonstrating the existence of both genotypes in natural P. bahamense sub-populations, as well as sxtA4 presence in P. bahamense from PR. These results provide insights on underlying genetic factors influencing the potential for toxin variability among natural sub-populations of HAB species and highlight the need to study the genetic diversity within HAB sub-populations at a fine level in order to identify the molecular mechanisms driving HAB evolution.

Diversity of dinoflagellate assemblages in coastal temperate and offshore tropical waters of Australia

BackgroundDinoflagellates are a ubiquitous and ecologically important component of marine phytoplankton communities, with particularly notable species including those associated with harmful algal blooms (HABs) and those that bioluminesce. High-throughput sequencing offers a novel approach compared to traditional microscopy for determining species assemblages and distributions of dinoflagellates, which are poorly known especially in Australian waters.ResultsWe assessed the composition of dinoflagellate assemblages in two Australian locations: coastal temperate Port Phillip Bay and offshore tropical waters of Davies Reef (Great Barrier Reef). These locations differ in certain environmental parameters reflecting latitude as well as possible anthropogenic influences. Molecular taxonomic assessment revealed more species than traditional microscopy, and it showed statistically significant differences in dinoflagellate assemblages between locations. Bioluminescent species and known associates of HABs were present at both sites. Dinoflagellates in both areas were mainly represented by the order Gymnodiniales (66%—82% of total sequence reads). In the warm waters of Davies Reef, Gymnodiniales were equally represented by the two superclades, Gymnodiniales sensu stricto (33%) and Gyrodinium (34%). In contrast, in cooler waters of Port Phillip Bay, Gymnodiniales was mainly represented by Gyrodinium (82%). In both locations, bioluminescent dinoflagellates represented up to 0.24% of the total sequence reads, with Protoperidinium the most abundant genus. HAB-related species, mainly represented by Gyrodinium, were more abundant in Port Phillip Bay (up to 47%) than at Davies Reef (28%), potentially reflecting anthropogenic influence from highly populated and industrial areas surrounding the bay. The entire assemblage of dinoflagellates, as well as the subsets of HAB and bioluminescent species, were strongly correlated with water quality parameters (R2 = 0.56–0.92). Significant predictors differed between the subsets: HAB assemblages were explained by salinity, temperature, dissolved oxygen, and total dissolved solids; whereas, bioluminescent assemblages were explained only by salinity and dissolved oxygen, and had greater variability.ConclusionHigh-throughput sequencing and genotyping revealed greater diversity of dinoflagellate assemblages than previously known in both subtropical and temperate Australian waters. Significant correlations of assemblage structure with environmental variables suggest the potential for explaining the distribution and composition of both HAB species and bioluminescent species.

Aquatic toxicity evaluations of PFOS and PFOA for five standard marine endpoints

Per- and poly-fluoroalkyl substances (PFAS) are emerging contaminants that are coming under increasing scrutiny. Currently, there is a paucity of effects data for marine aquatic life, limiting the assessment of ecological risks and compliance with water quality policies. In the present study, the toxicity of perfluorooctanesulfonic acid (PFOS) and perfluorooctanoic acid (PFOA) to four standard marine laboratory toxicity testing species, encompassing five endpoints, were evaluated: 1) 96-h embryo-larval normal development for the purple sea urchin (Strongylocentrotus purpuratus); 2) 48-h embryo-larval normal development and normal survival for the Mediterranean mussel (Mytilus galloprovincialis); 3) 96-h survival of opossum shrimp (Americamysis bahia); and 4) 24-h light output for the bioluminescent dinoflagellate Pyrocystis lunula. All species were tested using standard United States Environmental Protection Agency (USEPA) and/or American Society for Testing and Materials (ASTM) International protocols. For PFOS and PFOA, the order of species sensitivity, starting with the most sensitive, was M. galloprovincialis, S. purpuratus, P. lunula, and A. bahia. The range of median lethal or median effect concentrations for PFOS (1.1–5.1 mg L−1) and PFOA (10–24 mg L−1) are comparable to the relatively few toxicity effect values available for marine species. In addition to providing effects data for PFOA and PFOS, this study indicates these species and endpoints are sensitive to PFAS such that their use will be appropriate for deriving toxicity data with other PFAS in marine ecosystems.

Shear Stress‐Responsive Polymersome Nanoreactors Inspired by the Marine Bioluminescence of Dinoflagellates

AbstractSome marine plankton called dinoflagellates emit light in response to the movement of surrounding water, resulting in a phenomenon called milky seas or sea sparkle. The underlying concept, a shear‐stress induced permeabilisation of biocatalytic reaction compartments, is transferred to polymer‐based nanoreactors. Amphiphilic block copolymers that carry nucleobases in their hydrophobic block are self‐assembled into polymersomes. The membrane of the vesicles can be transiently switched between an impermeable and a semipermeable state by shear forces occurring in flow or during turbulent mixing of polymersome dispersions. Nucleobase pairs in the hydrophobic leaflet separate when mechanical force is applied, exposing their hydrogen bonding motifs and therefore making the membrane less hydrophobic and more permeable for water soluble compounds. This polarity switch is used to release payload of the polymersomes on demand, and to activate biocatalytic reactions in the interior of the polymersomes.

Shear Stress-Responsive Polymersome Nanoreactors Inspired by the Marine Bioluminescence of Dinoflagellates.

Some marine plankton called dinoflagellates emit light in response to the movement of surrounding water, resulting in a phenomenon called milky seas or sea sparkle. The underlying concept, a shear‐stress induced permeabilisation of biocatalytic reaction compartments, is transferred to polymer‐based nanoreactors. Amphiphilic block copolymers that carry nucleobases in their hydrophobic block are self‐assembled into polymersomes. The membrane of the vesicles can be transiently switched between an impermeable and a semipermeable state by shear forces occurring in flow or during turbulent mixing of polymersome dispersions. Nucleobase pairs in the hydrophobic leaflet separate when mechanical force is applied, exposing their hydrogen bonding motifs and therefore making the membrane less hydrophobic and more permeable for water soluble compounds. This polarity switch is used to release payload of the polymersomes on demand, and to activate biocatalytic reactions in the interior of the polymersomes.