Ecology

We are pleased to announce the winners of Rising Stars in Polymer Science 2022 as young influential. Polymer Journal has been enriched by the complex of wonderfully talented and diverse groups of these young scholars in addition to outstanding teams of well-established senior researchers. They bring a variety of new insights, both personal and professional, to the task of better understanding polymer science and engineering. Here they provide us with an array of novel observations drawn from such disciplines as synthesis, structure and physical properties and functions and applications. We believe our readers will appreciate the opportunity to learn new voices in this special issue.
Daisuke Aoki

Chiba University
Daisuke Aoki currently serves as an Associate Professor in the Department of Applied Chemistry and Biotechnology, Faculty of Engineering, at Chiba University. He obtained his Ph.D. from Tokyo Institute of Technology in 2014 under the tutelage of Prof. T. Takata. Between 2014 and 2017, he served as a specially appointed Assistant Professor in the group of Prof. T. Takata. From 2017 to 2022, he was an assistant professor at Tokyo Institute of Technology in the group of Prof. H. Otsuka. From 2018 to 2022, he also served as Japan Science and Technology Agency (JST) PRESTO Researcher. In 2022, he was appointed to his current position at Chiba University. His research is focused on the functional polymers with applications in materials science, the topological polymers, and the polymer recycling system. He has received the Award for Encouragement of Research in Polymer Science (2017) and The Young Scientist Lecture Award of the Kansai Regional Chapter (2020) from the Society of Polymer Science, Japan.
Rajashekar Badam

Japan Advanced Institute of Science and Technology
Rajashekar Badam completed M.Sc in Chemistry from Sri Sathya Sai Institute of Higher Learning, India in 2011. He received his Ph.D. in Materials Science from Japan Advanced Institute of Science and Technology (JAIST) with an “outstanding graduate award for the year 2016” in the area of carbon based electrocatalysis. Further he worked at Toyota Technological Institute as Postdoctoral fellow. In April 2018 he joined Matsumi lab, JAIST as Asst. Professor and since Oct 2020 he has been promoted to Sr. Lecturer in the same group. He has around 25 international publications and 10 patents (granted/pending) to his credit. His key research interest lies in organic-inorganic hybrid energy materials as catalysts, cathode material for metal air batteries, anode materials for Li-ion batteries and polymer binder materials for battery application.
Yu-Cheng Chiu

National Taiwan University of Science and Technology
Yu-Cheng Chiu joined the Department of Chemical Engineering at National Taiwan University of Science and Technology (Taiwan Tech). as a tenure-track assistant professor since August 2017. Currently, his major interests are the elastic and self-healing semiconducting materials, soft organic devices including transistor and transistor memory, and morphology characterization by synchrotron technique. Prior to joining the faculty, Yu-Cheng was a postdoc in the Zhenan Bao research group at Stanford University when he devoted on the research of intrinsically stretchable/healable semiconducting polymer and high-performance OFET by solution shearing technique. Before moving to Stanford, he received his Ph.D. degree under the supervision of Prof. Wen-Chang Chen in December 2012 from the Chem. E at National Taiwan University and then stayed in the same group for his first postdoctoral research until Oct. 2014. He also experienced international internship program as a Ph.D. student in 2010 and special appointed assistant professor position in 2018 for polymerization research in the group of Prof. Toyoji Kakuchi and Prof. Toshifumi Satoh at Hokkaido University.
Nagoya University
Yuya Doi received his Ph.D. degree under the supervision of Prof. Yushu Matsushita and Assoc. Prof. Atsushi Takano from Nagoya University in 2016. He worked as a Program-Specific Assistant Professor in the group of Prof. Hiroshi Watanabe at Kyoto University in 2016–2017, and was a visiting scholar in the group of Prof. Dimitris Vlassopoulos at FORTH, Greece in 2017. Then, he worked as a postdoctoral researcher at Nagoya University (in the group of Prof. Yushu Matsushita) from 2018, and at Forschungszentrum Jülich, Germany (in the group of Prof. Stephan Förster) from 2019. Since 2020, he has been an Assistant Professor at Nagoya University working with Prof. Yuichi Masubuchi and Assoc. Prof. Takashi Uneyama. His research interest is fundamental physical properties of model polymers studied by rheological and scattering methods.
Yuuka Fukui

Keio University
Yuuka Fukui received Ph.D. degree from Keio University in 2012 under the supervision of Professor Keiji Fujimoto. She was a JSPS research fellow (DC2) from 2010 to 2012. She joined the laboratory of Professor Keiji Fujimoto at Keio university as a research associate in 2012 and was promoted to an assistant professor in 2017. Her research interests focus on the design and synthesis of polymeric materials (particles, porous materials, membranes) and organic–inorganic hybrid materials inspired from biological systems. Her current research also includes development of functional materials to aim for applications in drug and cosmetic delivery systems and tissue engineering.
Mikihiro Hayashi

Nagoya institute of technology
Mikihiro Hayashi received his Ph.D. degree from Nagoya University (Prof. Yushu Matsushita group) in 2015. During his doctor course, he had been selected as a JSPS research fellow (DC2) and experienced researches in ESPCI Paris-Tech (Prof. Ludwik Leibler) and in Shanghai Jiao Tong University (Prof. Xinyuan Zhu). He then re-joined Ludwik Leibler’s group as a postdoc, and experienced another postdoc in Prof. Masatoshi Tokita in Tokyo institute of technology. In 2017, he became an assistant professor in Prof. Akinori Takasu group (Nagoya institute of technology), and currently manages his own laboratory as a PI. His research interest is the design of functional cross-linked materials. As recent awards, he won the SPSJ polymer research encouragement award (year—2019) and SPSJ award for the outstanding paper in Polymer Journal sponsored by ZEON (year—2021).
Kanazawa University
Asae Ito is an assistant professor under the Koh-hei Nitta’s laboratory; Polymer Physics Laboratory. She has received her B.S. in Chemistry in Tokyo University of Science in 2010, and M.S. in Tokyo Institute of Technology in 2012. She joined in R&D section of SHARP corporation and engaged in the fabrication of OLED devices (2012–2016). Then, she went on to Japan Advanced Institute of Science and Technology (JAIST) and obtained Ph.D. under the supervision of Prof. M. Yamaguchi in 2019 on polymer rheology. Her major interests are the correlation between structure and mechanical properties in glassy as well as semicrystalline polymeric materials.
Tomohiro Miyata

Tohoku University
Tomohiro Miyata received his B.S. in 2013 and Ph.D. in 2018 from the University of Tokyo. After working as a JSPS postdoctoral researcher at Tohoku University, he got a post of Assistant Professor at Tohoku University in 2019. He received several awards, including Young Scientist Award from the Japanese Society of Polymer Science and Dean’s Award FY2017 for the Best Doctoral Student from the School of Engineering, the University of Tokyo. He has worked on ceramics and liquid analysis using TEM techniques since 2013, and engaged in atomic- and nano-scale analysis on polymeric materials since 2018 in Jinnai group at Tohoku University.
Yuta Nishina

Okayama University
Yuta Nishina obtained his Ph.D. degree in Engineering from Okayama University in 2010. Then, he became an independent assistant professor at Research Core for Interdisciplinary Sciences, Okayama University, and was promoted to associate professor in 2014 and research professor in 2018. He has also been appointed as visiting professor at Florida State University (2011), Nanyang Technological University (2011–2012), University of Strasbourg (2017), and Osaka University (2017–2020). His research activities include JST PRESTO (2013–2017), JST CREST (2018—present and 2020—present), and Adjunct Professor at University of New England. He is currently working in multi-discipline research based on organic chemistry, such as nanocarbon production and functionalization, biomedicals, catalysis, and energy-related devices.
Yasunari Tamai

Kyoto University
Yasunari Tamai received his PhD from Kyoto University in 2013 on the excited state dynamics in nanostructured polymer systems. He joined the Optoelectronics group at the University of Cambridge as a postdoctoral fellow under the supervision of Prof Sir Richard Friend, where he focused on ultrafast charge separation at organic semiconductor heterojunctions. Since 2016, he has been an Assistant Professor at Kyoto University. From 2018 to 2022, he was also a JST PRESTO researcher. His current research interests include exciton and charge dynamics in organic semiconductors, particularly conjugated polymers.
Nanjing University
Ye Zhang is currently an associate professor at the College of Engineering and Applied Sciences at the Nanjing University. She received her Ph.D. degree in Macromolecular Chemistry and Physics from the Fudan University in 2018 and then joined the Harvard Medical School as a postdoctoral research fellow. Her research focuses on the development of soft electronics including batteries, sensors, and bioelectronic devices.
Tohoku University
Huie Zhu is an assistant professor in Graduate School of Engineering, Tohoku University. She received her B.Eng. (2008) and M.Eng. degrees (2011) from Zhengzhou University, China. Then, she obtained her Ph.D. degree in Applied Chemistry from Tohoku University in 2014 under the supervision of Prof. Masaya Mitsuishi. After that, she worked shortly as a postdoctoral researcher with Prof. Masaya Mitsuishi in Institute of Multidisciplinary Research for Advanced Materials (IMRAM), Tohoku University until 2015 and then became an assistant professor in the same institute. From 2020, she started her current position. Her research interests are development of siloxane-based hybrid polymer materials under mild conditions for various applications such as adhesives and thermally stable coatings and nanostructure control of ferroelectric polymers at interfaces for improved performance. She has received several awards from academic organizations and conference committees, such as the Promotion and Nurturing of Female Researchers Contribution Award from the Japan Society of Applied Physics (2019) and the Award for Encouragement of Research in Polymer Science from The Society of Polymer Science, Japan (2020).
Zhejiang Sci-Tech University
Biao Zuo received all his degrees from Zhejiang Sci-Tech University (Hangzhou, China); Chemistry (BSc, 2008), Physical Chemistry of Polymers (MSc, 2011) and Textile Materials (PhD, 2014). After completing the Ph.D. degree, he took a lecturer position at the Department of Chemistry, ZSTU. In 2017 and 2021, he was promoted to associated professor and full professor, respectively. He has worked for a while at Princeton University (2018–2020) and Kyushu University (2016) as a visiting scholar. He is also a principal investigator (PI) at Key Laboratory of Surface & Interface Science of Polymer Materials (SISPM) of Zhejiang Province. His research focuses mainly on molecular dynamics, glass transition, viscoelastic relaxation, rheology and tribology of polymers at surface, interface and under confinement, e.g., ultra-thin films. He has been awarded Chinese Chemical Society (CCS) Young Chemist Award (2021) for the contribution of “Revealing molecular mechanisms of polymer dynamics at surfaces and interfaces”. He is also a recipient of Excellent Young Investigator of NSFC (2021). More

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Here we assessed how species and functional diversity components of wild bee assemblages responded to increasing urbanization levels, using a large dataset encompassing recent surveys gathering 838 sampling sites located in natural, semi-natural and urban habitats of France, Belgium and Switzerland.We found a weak, but significant negative effect of the proportion of impervious surfaces in a 500 m radius around each site on local species richness of bee communities. Thus, sites with high soil sealing tended to host less species than those with low soil sealing. However, this trend was not observed when using human population density as an urbanization metric: sites with denser human populations hosted on average the same number of species as less densely populated sites.Concerning taxonomic homogenization of communities, we did not record any effects of urbanization, both in terms of impervious surfaces or human population density.Analyses of occurrence rates of bee functional traits revealed significant differences between poorly and highly urbanized communities, for both urbanization metrics. With higher human population density, probabilities of occurrence of above-ground nesters, generalist and small species increased, and a higher probability of occurrence of above-ground nesters, generalists and social bees were recorded in areas with high soil sealing.Therefore, we found overall consistent results linking urbanization and wild bees taxonomic as well as functional trait diversity, even though analyses stemmed from a combination of many independent studies covering a broad range of anthropized and natural aeras from western Europe. This further highlights the greater generalizability of those ecological trends throughout European temperate biomes compared to other studies typically focusing on a single city and its immediate vicinity.Two complementary metrics of urbanization intensityTo quantify urbanization, we used two variables: soil sealing12,16,19,36 in a 500 m radius, and the mean human population density, also in a 500 m radius, the latter variable being used only recently to assess pollinator responses to urban environments37,38. These two variables return different but complementary information concerning urban environments. Indeed, if soil sealing gives an idea as to how human activities impact land use, human population density helps distinguish between very dense urban areas and very impervious areas with lower densities of buildings. High human population density areas are usually associated with high levels of soil sealing, but the contrary is not true. Similarly, areas with low soil sealing are usually associated with low human population densities, but again, the opposite is not always true. Therefore, we found it informative to consider both variables when analyzing the response of wild bee assemblages to urbanization.Note that some specific habitat types, for example business districts, are exceptions to the rule. These places are indeed very densely urbanized, but with very low population density. However, no inventories have been carried out in these places, and thus will not be a problem for our study.Response of bee community species richness to urbanizationOne of our goals was to position this study in the context of the contrasting findings on pollinator communities and urbanization. Whereas no consistent trend is reported in literature15, our large dataset reveals that high soil sealing is detrimental to wild bee species richness. This offers a unified view of a trend that has been unequally evidenced from studies focusing on a single or few cities only. High proportions of soil sealing reduce the availability of nesting sites for ground-nesting bee species. This may in turn lower the species diversity of local assemblages, by filtering out ground-nesting bees, leaving mainly cavity-nesting bees. Furthermore, high levels of soil sealing can lead to depletion of floral resources, of extreme importance for bees, especially in highly disturbed environments such as cities39,40. Note that several previous studies report the opposite, with high local species richness of wild bees in urbanized habitats. However, these positive effects are often associated with intermediate levels of urbanization15,16, where private gardens and other green spaces may supply abundant floral resources, in conjunction with intermediate levels of soil sealing16,17,18,19,20,24.On the contrary, there was no significant relationship between local species richness and human population density. Recently, two recent studies have used this metric to analyze how urbanization impacts local diversity of bee, hoverfly37 or butterfly38 assemblages, and both studies report negative impacts of human population density. However, high levels of human population density do not necessarily correlate with low availability of floral resources or nesting sites for pollinating insects. Several studies show that densely-populated urban environments may be adequate habitats for pollinating insects, due to alternative management practices of urban green space41 and the year-round availability of ornamental flowers42,43. Here, the absence of a clear effect of human population density on local bee species richness masks a change in the species composition of the communities, as shown by the increasing proportion of cavity nesters, compared with ground nesters. Indeed, despite the lower availability of nesting resources for ground-nesters, cavity-nesters take over in high-density areas, where more concrete structures and buildings are present15, thus they may compensate for the loss of ground-nesting bee species.Wild bee community homogenization and urbanizationWe did not observe any relationship between mean pairwise β-diversity and the two metrics of urbanization. This result contrasts with those of Banaszak-Cibicka and Żmihorski (2020)44 who found more homogeneous wild bee communities in urban environments compared to non-urban ones. Similar results have been reported for bees, with homogenization of urban pollinator communities compared to rural ones28,45. Biotic homogenization in urban environments has also been reported for other taxa, for example birds46.In our study, when considering urbanization levels, either in terms of soil sealing or human population density, urban wild bee communities are not more or less taxonomically homogeneous than non-urban ones. It is important to note that this result does not imply that urban and non-urban wild bee communities are similar, but that the homogenization of wild bee communities is constant throughout the urbanization gradient. In other words, urban communities are as dissimilar as non-urban ones. Here, the β diversity values are quite high (ranging from 0.68 to 0.96), emphasizing that even urban areas have quite dissimilar communities when compared to each other. This high level of dissimilarity among wild bee communities in urban environments can be explained by the large range of biogeographical regions encompassed in our dataset (Fig. 5), as each of these regions harbors a specific wild bee fauna34.Local factors in cities might also explain these high levels of dissimilarity. We know for example that green space connectivity has effects on species richness, with more wild bee species and abundance in cities with more connected green spaces47. Another local explanation might come from contrasting green space management practices among cities. Not all cities have the same policies, and urban green space management is crucial to the establishment and sustainability of diverse pollinator communities14,15,48. Thus, we expect more dissimilar wild bee communities among cities with differing green space layout and management.Figure 5Grouped sampling sites (n = 532) in France, Belgium and Switzerland, with the biogeographical regions. In total, 238 sites belong to the Continental region, 178 to the Atlantic, 106 to de Mediterranean and 10 to the Alpine. This figure was generated using QGIS software, v3.10.13 (https://www.qgis.org/).Full size imageFunctional responses of bee communities to urbanizationSeveral studies have already shown trends on how urban areas filter wild bee communities based on their functional traits (see30 and49 for reviews). However, as for taxonomic diversity, it is often difficult to identify clear variation patterns50. Using our large dataset, we could identify typical wild bee functional traits that are favored in urban environments, thus informing on the average functional profiles of wild bee species that may thrive in cities. We found urban wild bees in general to be typically above-ground nesters and generalists, while different trends were established for their body size and sociality, depending on the considered urbanization metric (Fig. 6).Figure 6Summary picture of an urban bee community, compared to a non-urban one. This figure was generated using Inkscape v1.2 (https://inkscape.org/).Full size imageNesting habitsAbove-ground nesting species were more frequent with increasing urbanization than below-ground nesting ones, and this result was recorded with both urbanization metrics.This result is consistent with what was previously reported in the literature16,49,51,52. Indeed, cities, with high proportions of impervious surfaces and buildings, offer fewer nesting habitats to ground-nesting species15, nesting sites becoming a limiting factor39. On the other hand, above-ground nesters can do well in cities with the presence of man-made structures, depending on their ability to use them and on their availability53.The presence of green areas in cities can help ground-nesting bee species by offering more nesting opportunities and resources17. Several studies highlight the importance of parks and gardens in supporting bee biodiversity in cities12,18,31,54, which otherwise are constraining environments due to soil sealing.DietGeneralist species were more frequent in more urbanized sites than specialist ones, and this was recorded for both urbanization metrics.This is in accordance with what was previously found in the literature32,50,51,52,54,55, as specialist bee species depend on the presence of their host plants to complete their life-cycle, which are often scarce due to the rarefaction of native flowering resources. As one can find many exotic flowers in cities, especially in residential gardens and urban parks56, we expect to detect less oligolectic bee species in densely urbanized habitats57.Notwithstanding, Banaszak-Cibicka et al. (2018)20 found more oligolectic species in urban parks of Poznań (Poland) compared to a national park. Thus, urban areas are not always depleted of specialist species, and well-managed parks with preserved native floral resources can obviously support specialist wild bee species in cities58.Additionally, it is important to emphasize that the presence of an exotic plant species may concomitantly support an associated specialist bee species. In Poland, for instance, the spread of Bryonia dioica in urban environments also brought the Andrena florea wild bee species, specialized on this plant59.Body sizeWe recorded contrasting effects of the two urbanization metrics on wild bee body size: small species were more frequent in relation to higher human population density compared to large species, but we found no difference with the proportion of impervious surfaces. Contrasting impacts of urbanization on bee body size are also reported in the literature, with some studies finding little to no effect32,50, and some finding that urbanization often favors smaller bee species12,30,60. Bee body size is of particular importance because it is related to the foraging range of individuals61,62. In fragmented habitats, such as dense urban environments, distances between suitable nesting and feeding habitats may select for smaller species that can remain on small green spaces and rarely need to commute across several green spaces. Furthermore, small bees may be favored given that they need fewer floral resources than large bees, even though large bees can fly further62.This might also explain the difference in the response of bee body size to the two urbanization metric results. In densely populated cities, it is harder to fly between suitable habitats, even for larger bees, as higher buildings and structures may act as barriers to their movement. Indeed, it has been recently shown that the 3D structure of cities impacts wild bee community composition63. Thus, being able to fly further might no longer be an advantage, and larger bees, requiring more floral resources than smaller ones, might be selected against. On the contrary, very impervious areas do not always host high building density (for example, as in the case of parking lots), thus making it easier for large wild bees to fly between bare soil areas.Densely populated areas might also exhibit warmer temperatures due to the urban heat island effect, and this could, in turn, result in the selection of smaller individuals, as we know that in cities, higher temperature results in smaller body sizes64.SocialityWe also recorded contrasting effects of the two urbanization metrics on sociality: social species were more frequent in relation to higher proportion of impervious surface compared to solitary ones, but no effect was recorded with human population density. This is in agreement with a recent literature review that reports on no consensus concerning the response of this trait to urbanization30.However, some urban habitats are shown to host more social species than rural habitats20,32, which may be linked to better reproductive success in cities compared to rural habitats such as agricultural environments65, an explanation that is consistent with our results on the soil sealing—sociality relationship.Conclusion, limits & future directionsOverall, our findings suggest that urban environment filters wild bee communities based on their functional traits. Our results also underscore different impacts of urbanization metrics on local species diversity, with a significant negative impact of soil sealing. On the contrary, both soil sealing and human population densities create strong functional filtering of trait assemblages.These results are particularly relevant since they arise from a range of independent studies, thus providing a general view on the wild bee communities in urban environments from western Europe. Since this study covers different biogeographical zones, it further underlines its applicability to other temperate countries. We therefore expect similar patterns to shape wild bee communities in urbanized areas from other temperate regions, but further confirmatory studies would be welcome.Our study also delivers a clear message concerning wild bee communities in urban environments. Urban environments cannot compare with non-urban ones in terms of species richness and trait diversities of bee communities. However, simple management practices of urban green spaces, such as differentiated management, or simply low management66, may help in maintaining this diversity. Indeed, not all green spaces are equally valuable in supporting wild bees, and pollinator assemblages in general49. For example, it has been shown that pollinator richness was positively influenced by green space size, but also by management measures such as mowing67. Increasing the quantity of floral resources and their spatio-temporal availability and diversity40,68 could also help conserving pollinator communities and pollination function in cities69, as long as these resources are native or attractive to pollinators.We can then hypothesize that changes in managing practices could help increase functional diversity of bees in cities, with specialist and ground-nesting species being found more frequently in these low-managed urban areas.Finally, if managing urban green space is of great importance to protect biodiversity in cities, it is crucial to involve all stakeholders, especially residents70 to achieve efficient and socially-accepted measures.In the future, it will be important to consider intra-city landscape variation, and see how urban characteristics might influence taxonomic and trait diversity. This will surely allow us to better understand the dynamics shaping wild bee communities in urban environments. More

Measurement of colony strength traits and Varroa destructor loadHives located in the natural area (henceforth natural hives or colonies) weighed the most (hive weight, Kruskal–Wallis test, p  More

All Alteromonas strains support long-term survival of Prochlorococcus under N starvationPrevious research showed that Prochlorococcus, and to some extent Synechococcus depend on co-occurring heterotrophic bacteria to survive various types of stress, including nitrogen starvation [33, 34, 42, 43]. At the first encounter between previously axenic Prochlorococcus and Alteromonas (E1), all co-cultures and axenic controls grew exponentially (Fig. 1B, C). However, all axenic cultures showed a rapid and mostly monotonic decrease in fluorescence starting shortly after the cultures stopped growing, reaching levels below the limit of detection after ~20–30 days. None of the axenic Prochlorococcus cultures were able to re-grow when transferred into fresh media after 60 days (Fig. 1C). In contrast, the decline of co-cultures rapidly slowed, and in some cases was interrupted by an extended “plateau” or second growth stage (Fig. 1B). Across multiple experiments, 92% of the co-cultures contained living Prochlorococcus cells for at least 140 days, meaning that they could be revived by transfer into fresh media. Thus, the ability of Alteromonas to support long-term N starvation in Prochlorococcus was conserved in all analyzed strains.Fig. 1: Experimental designs and overview of the dynamics of Prochlorococcus-Alteromonas co-cultures from first encounter and over multiple transfers.A Schematic illustration of the experimental design. One ml from Experiment E1 was transferred into 20 ml fresh media after 100 days, starting experiment E2. Experiment E2 was similarly transferred into fresh media after 140 days, starting experiment E3. Additional experiments replicating these transfers are described in Supplementary Fig. S1. B Overview of the growth curves of the 25 Prochlorococcus-Alteromonas co-cultures over three transfers spanning ~1.2 years (E1, E2 and E3). Results show mean + standard error from biological triplicates, colored by Prochlorococcus strain as in panel D. C Axenic Prochlorococcus grew exponentially in E1 but failed to grow when transferred into fresh media after 60, 100, or 140 days. Axenic Alteromonas cultures were counted after 60 and 100 days, as their growth cannot be monitored sensitively and non-invasively using fluorescence (optical density is low at these cell numbers). D High reproducibility and strain-specific dynamics of the initial contact between Prochlorococcus and Alteromonas strains (E1). Three biological replicates for each mono-culture and co-culture are shown. Note that the Y axis is linear in panels B, C and logarithmic in panel D. Au: arbitrary units.Full size imageIt has previously been shown that Prochlorococcus MIT9313 is initially inhibited by co-culture with Alteromonas HOT1A3, while Prochlorococcus MED4 is not [12, 32]. This “delayed growth” phenotype was observed here too, was specific to MIT9313 (not observed in other Prochlorococcus strains) and occurred with all Alteromonas strains tested (Fig. 1D). MIT9313 belongs to the low-light adapted clade IV (LLIV), which are relatively distant from other Prochlorococcus strains and differ from them in multiple physiological aspects including the structure of their cell wall [44], the use of different (and nitrogen-containing) compatible solutes [45], and the production of multiple peptide secondary metabolites (lanthipeptides, [46, 47]). LLIV cells also have larger genomes, and are predicted to take up a higher diversity of organic compounds such as sugars and amino acids [48]. It is intriguing that specifically this strain, which has higher predicted metabolic and regulatory flexibilities [49], is the only one initially inhibited in co-culture with Alteromonas.Differences in co-culture phenotype are related to Prochlorococcus and not Alteromonas strains and occur primarily during the decline stageWhile co-culture with all Alteromonas strains had a major effect on Prochlorococcus viability after long-term starvation, there was no significant effect of co-culture on traditional metrics of growth such as maximal growth rate, maximal fluorescence, and lag phase (with the exception of the previously described inhibition of MIT9313; Fig. 2A–C). However, a visual inspection of the growth curves suggested subtle yet consistent differences in the shape of the growth curve, and especially the decline phase, between the different Prochlorococcus strains in the co-cultures (Fig. 1D). To test this, we used the growth curves as input for a principal component analysis (PCA), revealing that the growth curves from each Prochlorococcus strain clustered together, regardless of which Alteromonas strain they were co-cultured with (Fig. 2D). The growth curves of all high-light adapted strains (MED4, MIT9312, and MIT0604) were relatively similar, the low-light I strain NATL2A was somewhat distinct, and the low-light IV strain MIT9313 was a clear outlier (Fig. 2D), consistent with this strain being the only one initially inhibited in all co-cultures. Random forest classification supported the observation that the growth curve shapes were affected more by the Prochlorococcus rather than Alteromonas strains, and also confirmed the visual observation that most of the features differentiating between Prochlorococcus strains occurred during culture decline (random forest is a supervised machine learning algorithm explained in more detail in Supplementary Text S2; see also Supplementary Fig. S2). Thus, co-culture with Alteromonas affects the decline stage of Prochlorococcus in co-culture in a way that differs between Prochlorococcus but not Alteromonas strains.Fig. 2: Growth analysis and principal component analysis (PCA) of the growth curves from all co-cultures during 140 days of E1.A Growth rate, B Maximum fluorescence, and C duration of lag phase during experiment E1. Box-plots represent mean and 75th percentile of co-cultures, circles represent measurements of individual cultures of the axenic controls. The only significant difference between axenic and co-cultures is in the length of the lag phase for MIT9313 (Bonferroni corrected ANOVA, p  More