The importance of the vertical structure of vegetation cover for the functioning, management and conservation of ecosystems has received particular attention from ecologists in the last decades. Canopy architecture has many implications for light extinction coefficient, temperature variation reduction, self-shading which are all key parameters for the structuring and functioning of different ecosystems such as grasslands [1,2], forests [3,4], phytoplankton communities [5, 6], macroalgal populations [7] and even underwater animal forests such as octocoral communities [8].

This research topic, therefore, benefits from a large body of literature and the facilitative role of self-shadowing is no longer in question. However, it is always puzzling to note that some of the most common ecosystems turn out to be amongst the least known. This is precisely the case of the Fucus serratus communities which are widespread in Northeast Atlantic along the Atlantic coast of Europe from Svalbard to Portugal, as well as Northwest Atlantic & Gulf of St. Lawrence, easily accessible at low tide, but which have comparatively received less attention than more emblematic macro-algal communities such as Laminariales.

The lack of attention paid to these most common Fucales is particularly critical as some species such as F. serratus are proving to be particularly vulnerable to environmental change, leading to a predicted northward retreat from its current southern boundary [9].

In the present study [10], the authors showed the importance of the vegetation cover in resisting tide-induced environmental stresses. The canopy of F. serratus mitigates stress levels experienced in the lower layers during emersion, while various acclimation strategies take over to maintain the photosynthetic apparatus in optimal conditions.

They hereby highlight adaptation mechanisms to the extreme environment represented by the intertidal zone. These adaptation strategies were expected and similar mechanisms had been shown at the cellular level previously [11]. The earliest studies on the subject have shown that the structure of the bottom, the movement of water, and light availability all "influence the distribution of Fucaceae and disturb the regularity of their fine zonation, which itself is caused by the most important factor, desiccation", as Zaneveld states in his review [12]. He observed that the causes of the zonal distribution of marine algae are numerous, and identified several points of interest such as the relative period of emersion, the rapidity of desiccation, the loss of water, and the thickness of the cell walls.

The present study thus highlights the existence of facilitative mechanisms associated with F. serratus canopy and nicely confirms previous work with in situ observations. It also highlights the importance of the vegetative cover in combating desiccation and introduces the dampening effect as a facilitating mechanism.

The effect of the vegetation cover can sometimes even be felt beyond its immediate area of influence. A recent study shows that ground-level ozone is significantly reduced by the combined effects of canopy shading and turbulence [4]. Below the canopy, the light intensity becomes sufficiently low which inhibits ozone formation due to the decrease in the rates of hydroxyl radical formation and the rates of conversion of nitrogen dioxide to nitrogen oxide by photolysis. In addition, reductions in light levels associated with foliage promote ozone-destroying reactions between plant-emitted species, such as nitric oxide and/or alkenes, and ozone itself. The reduction in diffusivity slows the upward transport of surface emitted species, partially decoupling the area under the canopy from the rest of the atmosphere.

By analogy with the work of Makar et al [4], and in the light of the results provided by the authors of this study, one may wonder whether the canopy dampening of F. serratus communities (and other common fucoids widely distributed on our coasts) might not also influence atmospheric chemistry, both at the Earth's surface and in the atmospheric boundary layer. The lack of accumulation of reactive oxygen species under the canopy found by the authors is consistent with this hypothesis and suggests that the damping effect of F. serratus may well have much wider consequences than expected.

References

[1] Jurik TW, Kliebenstein H (2000) Canopy Architecture, Light Extinction and Self-Shading of a Prairie Grass, Andropogon Gerardii. The American Midland Naturalist, 144, 51–65. http://www.jstor.org/stable/3083010

[2] Mitchley J, Willems JH (1995) Vertical canopy structure of Dutch chalk grasslands in relation to their management. Vegetatio, 117, 17–27. https://doi.org/10.1007/BF00033256

[3] Kane VR, Gillespie AR, McGaughey R, Lutz JA, Ceder K, Franklin JF (2008) Interpretation and topographic compensation of conifer canopy self-shadowing. Remote Sensing of Environment, 112, 3820–3832. https://doi.org/10.1016/j.rse.2008.06.001

[4] Makar PA, Staebler RM, Akingunola A, Zhang J, McLinden C, Kharol SK, Pabla B, Cheung P, Zheng Q (2017) The effects of forest canopy shading and turbulence on boundary layer ozone. Nature Communications, 8, 15243. https://doi.org/10.1038/ncomms15243

[5] Shigesada N, Okubo A (1981) Analysis of the self-shading effect on algal vertical distribution in natural waters. Journal of Mathematical Biology, 12, 311–326. https://doi.org/10.1007/BF00276919

[6] Barros MP, Pedersén M, Colepicolo P, Snoeijs P (2003) Self-shading protects phytoplankton communities against H2O2-induced oxidative damage. Aquatic Microbial Ecology, 30, 275–282. https://doi.org/10.3354/ame030275

[7] Ørberg SB, Krause-Jensen D, Mouritsen KN, Olesen B, Marbà N, Larsen MH, Blicher ME, Sejr MK (2018) Canopy-Forming Macroalgae Facilitate Recolonization of Sub-Arctic Intertidal Fauna and Reduce Temperature Extremes. Frontiers in Marine Science, 5. https://doi.org/10.3389/fmars.2018.00332

[8] Nelson H, Bramanti L (2020) From Trees to Octocorals: The Role of Self-Thinning and Shading in Underwater Animal Forests. In: Perspectives on the Marine Animal Forests of the World (eds Rossi S, Bramanti L), pp. 401–417. Springer International Publishing, Cham. https://doi.org/10.1007/978-3-030-57054-5_12

[9] Jueterbock A, Kollias S, Smolina I, Fernandes JMO, Coyer JA, Olsen JL, Hoarau G (2014) Thermal stress resistance of the brown alga Fucus serratus along the North-Atlantic coast: Acclimatization potential to climate change. Marine Genomics, 13, 27–36. https://doi.org/10.1016/j.margen.2013.12.008

[10] Migné A, Duong G, Menu D, Davoult D, Gévaert F (2021) Dynamics of Fucus serratus thallus photosynthesis and community primary production during emersion across seasons: canopy dampening and biochemical acclimation. HAL, hal-03079617, ver. 4 peer-reviewed and recommended by Peer community in Ecology. https://hal.archives-ouvertes.fr/hal-03079617

[11] Lichtenberg M, Kühl M (2015) Pronounced gradients of light, photosynthesis and O2 consumption in the tissue of the brown alga Fucus serratus. New Phytologist, 207, 559–569. https://doi.org/10.1111/nph.13396

[12] Zaneveld JS (1937) The Littoral Zonation of Some Fucaceae in Relation to Desiccation. Journal of Ecology, 25, 431–468. https://doi.org/10.2307/2256204

The concept of holobiont dates back to more than thirty years, it was primarily coined to hypothesize the importance of symbiotic associations to generate significant evolutionary novelties. Quickly adopted to describe the now well-studied system formed by zooxanthella associated corals, this concept expanded much further after the emergence of High-Throughput Sequencing and associated progresses in metabarcoding and metagenomics.
Holobionts – defined as the association between an individual host and its microbiota - are now increasingly described at sea and on land. The opinion article by Dittami et al. [1] provides a synthetic overview of marine holobionts. It retraces the history of the holobiont concept, recalls the main mechanisms underlying the association between hosts and microbial communities, highlights the influence of these symbioses on marine ecosystem functioning, and outlines current tools and future lines of research.
In particular, the article discusses some particularities of marine systems, such as the strong connectivity allowing an exchange of microorganisms and chemical signals between and within holobionts.
The authors advocate the need to bridge the gap between large scale exploration studies and smaller scale mechanistic studies, by conducting interdisciplinary research (combining physiology, biochemistry, ecology, experimentation and computational modeling) on some keystone holobionts.
Finally, one strength of the paper by Dittami et al. [1] is that it places the concept of the holobiont in an applied research framework. Several possible applications of knowledge on host-microbiota interactions are suggested, both in the field of aquaculture and that of monitoring the health of marine ecosystems. This article contains all the necessary elements for someone who would like to jump into the study of the holobionths in the marine world.

References
[1] Dittami SM, Arboleda E, Auguet J, Bigalke A, Briand E, Cardenas P, Cardini U, Decelle J, Engelen AH, Eveillard D, Gachon CMM, Griffiths SM, Harder T, Kayal E, Kazamia E, Lallier FH, Medina M, Marzinelli E, Morganti T, Núñez Pons L, Prado S, Pintado J, Saha M, Selosse M, Skillings D, Stock W, Sunagawa S, Toulza E, Vorobev A, Leblanc C, Not F. (2020). A community perspective on the concept of marine holobionts: current status, challenges, and future directions. Zenodo, ver. 4 peer-reviewed and recommended by PCI Ecology. doi: 10.5281/zenodo.3696771

Polygyny is assumed to have led to the evolution of strong sexual size dimorphism (SSD) in mammals, males often being heavier or showing more developed armaments than females (Weckerly 1998; Loison et al. 1999; Pérez‐Barbería et al. 2002).  SSD generally increases with the degree of polygyny of the species. However, the degree of SSD, and particularly of sexual mass dimorphism, is not fixed for each species, and differences exist between populations (Blanckenhorn et al. 2006; Cox & Calsbeek 2010) or even between seasons within populations (Rughetti & Festa‐Bianchet 2011).

In this study, Hewison et al. propose that studying seasonal variation in sexual mass dimorphism and how this can be affected by winter harshness and latitude allows us to better assess the energetic costs associated with the eco-evolutionary constraints acting on each sex. To achieve their goal, Hewison et al. use a formidable, long-term dataset of over 7,000 individuals, in five roe deer populations (Capreolus capreolus), from south-west France and Sweden.

According to the authors, sexual mass dimorphism should be at its lowest in early spring in this species due to a stronger trade-off between antler growth and body weight maintenance in males over winter than in females. Furthermore, harsher conditions, varying both in time and space (i.e., Sweden vs. France), should increase winter weight loss, and thus, mass change differences between the sexes should be stronger and show more variation in Sweden than in France.

Their results support their hypotheses. In the two Swedish populations, males lost more mass than females. In the three French populations, males maintain their body mass while females gain some over the winter. Because of these sex-dependent loss/gain in body mass, sexual dimorphism was stronger early in the winter and null at the onset of spring. Furthermore, sexual dimorphism was stronger in southern than in northern populations. In France, males weighed about 10% more than females, while they weighed about 5% more in Sweden. Roe deer, however, do not show any dimorphism early in the spring, when males start defending their territory. 

The authors also found more variation in mass change among years in Swedish than in French roe deer, suggesting a stronger effect of winter severity on the dynamics of mass change in northern than in southern populations. The authors interpret the decrease in sexual dimorphism throughout the winter by the fact that, during this period, the energetic cost paid by males associated with the growth of their antlers and the effort of establishing their mating territory. They thus attribute the greater mass change in males to the competitive allocation of resources to antler growth or body mass. They also discuss the low probability that such sex differences in mass change could be caused by females’ gestation in this species.

Interestingly, Hewison et al. found that individual differences represented more than 70% of the total variation in body mass, and the low estimated among-individual variance in slopes with time might indicate that, despite a lower SSD, selection pressures on body mass can still be maintained at times when body mass may play an important role, such as in spring with territorial defense or later during mating (Vanpé et al. 2010). 

I recommend this article because it produces strong results, which show, without a shadow of a doubt, sex differences in their seasonal mass changes, resulting in a marked seasonal variation in SSD. The differences observed between southern and northern populations confirm the idea that the severity of the winters endured by these populations acts as a constraint on the deer's patterns of mass change. I hope this study will encourage more examinations of how eco-evolutionary constraints affect the sexual size dimorphism.

References

Blanckenhorn, W. U., Stillwell, R. C., Young, K. A., Fox, C. W., & Ashton, K. G. (2006). When Rensch meets Bergmann: does sexual size dimorphism change systematically with latitude? Evolution, 60(10), 2004-2011. https://doi.org/10.1554/06-110.1

Cox, R. M., & Calsbeek, R. (2010). Sex-specific selection and intraspecific variation in sexual size dimorphism. Evolution, 64(3), 798-809. https://doi.org/10.1111/j.1558-5646.2009.00851.x

Hewison M, Bonnot N, Gaillard JM, Kjellander P, Lemaitre J-F, Morellet N. and Pellerin M (2024) Body mass change over winter is consistently sex-specific across roe deer (Capreolus capreolus) populations. bioRxiv, ver.4 peer-reviewed and recommended by PCI Ecology https://doi.org/10.1101/2022.09.09.507329

Loison, A., Gaillard, J. M., Pélabon, C., & Yoccoz, N. G. (1999). What factors shape sexual size dimorphism in ungulates? Evolutionary Ecology Research, 1(5), 611-633. https://www.evolutionary-ecology.com/issues/v01n05/jjar1019.pdf

Pérez‐Barbería, F. J., Gordon, I. J., & Pagel, M. (2002). The origins of sexual dimorphism in body size in ungulates. Evolution, 56(6), 1276-1285. https://doi.org/10.1111/j.0014-3820.2002.tb01438.x

Rughetti, M., & Festa‐Bianchet, M. (2011). Seasonal changes in sexual size dimorphism in northern chamois. Journal of Zoology, 284(4), 257-264. https://doi.org/10.1111/j.1469-7998.2011.00800.x

Vanpé, C., Gaillard, J. M., Kjellander, P., Liberg, O., Delorme, D., & Hewison, A. M. (2010). Assessing the intensity of sexual selection on male body mass and antler length in roe deer Capreolus capreolus: is bigger better in a weakly dimorphic species? Oikos, 119(9), 1484-1492. https://doi.org/10.1111/j.1600-0706.2010.18312.x

Weckerly, F. W. (1998). Sexual-size dimorphism: influence of mass and mating systems in the most dimorphic mammals. Journal of Mammalogy, 79(1), 33-52. https://doi.org/10.2307/1382840

Individuals ability to withstand abiotic and biotic stresses is crucial to the maintenance of populations at climate edge of tree species distribution. We start to have a detailed understanding of tree growth response and to a lesser extent mortality response in these populations. In contrast, our understanding of the response of tree fecundity and recruitment remains limited because of the difficulty to monitor it at the individual tree level in the field. Tree recruitment limitation is, however, crucial for tree population dynamics [1-2].
In their study Oddou-Muratorio et al. [3] use a new method that they recently developed that jointly estimate male and female effective fecundity in natural populations using naturally established seedlings [4]. Their method uses a spatially explicit Bayesian analysis based on molecular markers and parentage analyses (MEMM program [4]). They apply this method to an unmanaged Beech forest at the southern edge of Beech distribution, where tree defoliation – taken as an integrative indicator of tree abiotic and biotic stress – and growth have been monitored for all adult trees.
This allows the authors to explore alternative hypothesis about tree fecundity response to stress. In one hand, biotic and abiotic stresses are thought to negatively impact tree fecundity. In the other hand, management and studies of orchard fruit tree support the idea that stress could trigger a compensatory increase of fecundity at the cost of other performances such as growth and survival.
They show that both growth and female fecundity are negatively affected by defoliation. There was no evidence that stresses trigger a compensatory increase of fecundity. Yet, they also found that, for large highly defoliated trees, there was a trade-off between growth and female fecundity. Some individuals are able to mitigate stress impact on fecundity by decreasing their growth. It is difficult to understand with available data what is driving such divergent responses between defoliated individuals. This could be related to differences in micro-environmental conditions or genetic background of individual trees. Such individual-level difference in response to stress could be crucial to understand tree populations response to ongoing climate change. This study clearly opens exciting new perspectives to apply such new methods to understand the role of fecundity on tree population dynamics. For instance, could we apply this method across the species distribution to understand how effective fecundity and its response to abiotic stress change between southern edge populations, core populations, and northern edge populations? Using time-series retrieved from such analysis can we disentangle the effect of different climatic drivers? It would also be interesting to see how such results can contribute to analyses covering the full tree life cycle to understand the contribution of fecundity response to population and evolutionary.

References

[1] Clark, J. S. et al. (1999). Interpreting recruitment limitation in forests. American Journal of Botany, 86(1), 1-16. doi: 10.2307/2656950
[2] Petit, R. J., and Hampe, A. (2006). Some evolutionary consequences of being a tree. Annu. Rev. Ecol. Evol. Syst., 37, 187-214. doi: 10.1146/annurev.ecolsys.37.091305.110215
[3] Oddou-Muratorio, S., Petit, C., Journe, V., Lingrand, M., Magdalou, J. A., Hurson, C., Garrigue, J., Davi, H. and Magnanou, E. (2019). Crown defoliation decreases reproduction and wood growth in a marginal European beech population. bioRxiv, 474874, ver. 4 peer-reviewed and recommended by PCI Ecology. doi: 10.1101/474874
[4] Oddou‐Muratorio, S. and Klein, E. K. (2008). Comparing direct vs. indirect estimates of gene flow within a population of a scattered tree species. Molecular Ecology, 17(11), 2743-2754. doi: 10.1111/j.1365-294X.2008.03783.x

"Good and evil are so close as to be chained together in the soul"
Robert Louis Stevenson, Dr. Jekyll and Mr. Hyde

Maternally inherited symbionts—microorganisms that pass from a female host to her progeny—have two main ways of increasing their own fitness. First, they can increase the fecundity or viability of infected females. This “positive fitness effects” strategy is the one commonly used by mutualistic symbionts, such as Buchnera aphidicola—the bacterial endosymbiont of the pea aphid, Acyrthosiphon pisum [4]. Second, maternally inherited symbionts can manipulate the reproduction of infected females in a way that enhances symbiont transmission at the expense of host fitness. A famous example of this “reproductive parasitism” strategy is the cytoplasmic incompatibility (CI) [3] induced by bacteria of the genus Wolbachia in their arthropod and nematode hosts. CI works as a toxin-antidote system, whereby the sperm of infected males is modified in a lethal way (toxin) that can only be reverted if the egg is also infected (antidote) [1]. As a result, CI imposes a kind of conditional sterility on their hosts: while infected females are compatible with both infected and uninfected males, uninfected females experience high offspring mortality if (and only if) they mate with infected males [7].

These two symbiont strategies (positive fitness effects versus reproductive parasitism) have been traditionally studied separately, both empirically and theoretically. However, it has become clear that the two strategies are not mutually exclusive, and that a reproductive parasite can simultaneously act as a mutualist—an infection type that has been dubbed “Jekyll and Hyde” [6], after the famous novella by Robert Louis Stevenson about kind scientist Dr. Jekyll and his evil alter ego, Mr. Hyde. In important previous work, Zug and Hammerstein [7] analyzed the consequences of positive fitness effects on the dynamics of different kind of infections, including “Jekyll and Hyde” infections characterized by CI and other reproductive parasitism strategies. Building on this and related modeling framework, Karisto et al. [2] re-investigate and expand on the interplay between positive fitness effects and reproductive parasitism in Wolbachia infections by focusing on CI in both diplodiploid and haplodiploid populations, and by paying particular attention to the mathematical assumption structure underlying their results.

Karisto et al. begin by reviewing classic models of Wolbachia infections in diplodiploid populations that assume a “negative fitness effect” (modeled as a fertility penalty on infected females), characteristic of a pure strategy of reproductive parasitism. Together with the positive frequency-dependent effects due to CI (whereby the fitness benefits to symbionts infecting females increase with the proportion of infected males in the population) this results in population dynamics characterized by two stable equilibria (the Wolbachia-free state and an interior equilibrium with a high frequency of Wolbachia-carrying hosts) separated by an unstable interior equilibrium. Wolbachia can then spread once the initial frequency is above a threshold or an invasion barrier, but is prevented from fixing by a proportion of infections failing to be passed on to offspring. Karisto et al. show that, given the assumption of negative fitness effects, the stable interior equilibrium can never feature a Wolbachia prevalence below one-half. Moreover, they convincingly argue that a prevalence greater than but close to one-half is difficult to maintain in the presence of stochastic fluctuations, as in these cases the high-prevalence stable equilibrium would be too close to the unstable equilibrium signposting the invasion barrier.

Karisto et al. then relax the assumption of negative fitness effects and allow for positive fitness effects (modeled as a fertility premium on infected females) in a diplodiploid population. They show that positive fitness effects may result in situations where the original invasion threshold is now absent, the bistable coexistence dynamics are transformed into purely co-existence dynamics, and Wolbachia symbionts can now invade when rare. Karisto et al. conclude that positive fitness effects provide a plausible and potentially testable explanation for the low frequencies of symbiont-carrying hosts that are sometimes observed in nature, which are difficult to reconcile with the assumption of negative fitness effects. 

Finally, Karisto et al. extend their analysis to haplodiploid host populations (where all fertilized eggs develop as females). Here, they investigate two types of cytoplasmic incompatibility: a female-killing effect, similar to the CI effect studied in diplodiploid populations (the “Leptopilina type” of Vavre et al. [5]) and a masculinization effect, where CI leads to the loss of paternal chromosomes and to the development of the offspring as a male (the “Nasonia type” of Vavre et al. [5]). The models are now two-sex, which precludes a complete analytical treatment, in particular regarding the stability of fixed points. Karisto et al. compensate by conducting large numerical analyses that support their claims. Importantly, all main conclusions regarding the interplay between positive fitness effects and reproductive parasitism continue to hold under haplodiploidy. 

All in all, the analysis and results by Karisto et al. suggest that it is not necessary to resort to classical (but depending on the situation, unlikely) mechanisms, such as ongoing invasion or source-sink dynamics, to explain arthropod populations featuring low-prevalent Wolbachia infections. Instead, low-frequency equilibria might be simply due to reproductive parasites conferring beneficial fitness effects, or Wolbachia symbionts playing Dr. Jekyll (positive fitness effects) and Mr. Hyde (cytoplasmatic incompatibility). 

References

[1] Beckmann JF, Bonneau M, Chen H, Hochstrasser M, Poinsot D, Merçot H, Weill M, Sicard M, Charlat S (2019) The Toxin–Antidote Model of Cytoplasmic Incompatibility: Genetics and Evolutionary Implications. Trends in Genetics, 35, 175–185. https://doi.org/10.1016/j.tig.2018.12.004

[2] Karisto P, Duplouy A, Vries C de, Kokko H (2022) Positive fitness effects help explain the broad range of Wolbachia prevalences in natural populations. bioRxiv, 2022.04.11.487824, ver. 5 peer-reviewed and recommended by Peer Community in Ecology. https://doi.org/10.1101/2022.04.11.487824

[3] Laven H (1956) Cytoplasmic Inheritance in Culex. Nature, 177, 141–142. https://doi.org/10.1038/177141a0

[4] Perreau J, Zhang B, Maeda GP, Kirkpatrick M, Moran NA (2021) Strong within-host selection in a maternally inherited obligate symbiont: Buchnera and aphids. Proceedings of the National Academy of Sciences, 118, e2102467118. https://doi.org/10.1073/pnas.2102467118

[5] Vavre F, Fleury F, Varaldi J, Fouillet P, Bouletreau M (2000) Evidence for Female Mortality in Wolbachia-Mediated Cytoplasmic Incompatibility in Haplodiploid Insects: Epidemiologic and Evolutionary Consequences. Evolution, 54, 191–200. https://doi.org/10.1111/j.0014-3820.2000.tb00019.x

[6] Zug R, Hammerstein P (2015) Bad guys turned nice? A critical assessment of Wolbachia mutualisms in arthropod hosts. Biological Reviews, 90, 89–111. https://doi.org/10.1111/brv.12098

[7] Zug R, Hammerstein P (2018) Evolution of reproductive parasites with direct fitness benefits. Heredity, 120, 266–281. https://doi.org/10.1038/s41437-017-0022-5