BRIEFING NOTE

DATE: 21/04/2021

Effects of the Chernobyl and Fukushima accidents on ecosystems:
current knowledge

In the mass media, the public was able to see both reports and works of fiction on the flora and faunapresent in areas contaminated by the major nuclear accidents which occurred at the Chernobyl Nuclear
Power Plant on 26 April 1986 and the Fukushima Daiichi Nuclear Power Plant on 11 March 2011.
The observations presented in these reports raise questions about the effects of the contamination ofthese sites on fauna and flora. There has been a lot of research on some wild species in these areas,
and this research is helping to better understand the effects of radioactive contamination on
ecosystems - radioactive contamination that has been present on these sites for 35 years at Chernobyl
and 10 years at Fukushima.

Introduction
The Chernobyl and Fukushima-Daiichi nuclear accidents resulted in significant radioactive releases into
the environment. The characteristics of these releases are described and compared in the brief:
Fukushima and Chernobyl accidents: Different impacts on the environment published on the IRSN
website.
The radionuclides emitted during these nuclear accidents were deposited, then dispersed andredistributed in the environment over the years. The flora and fauna in these areas are exposed toionizing radiation in two ways: external exposure induced by radiation emitted by radionuclides
present in the environment, and internal exposure induced by radionuclides ingested, inhaled, orabsorbed by wildlife.
The radionuclides emitted during the accidents led to acute exposure of wildlife immediately after theaccident, and this exposure had deleterious effects on certain species in the contaminated areas.
Today, the individuals constituting the wildlife in the contaminated areas remain chronically exposed
to radionuclides present in the environment every day and come from successive generations whichhave been produced by individuals that were initially exposed or by those which subsequently
migrated to contaminated areas.
These observations raise various questions, such as are the effects observed in the short term stillvisible? Have new effects, unrelated to short-term effects, appeared?
The work presented below aims to shed light on these questions and is based on recent knowledge(published mainly since 2016) on the effects on the wildlife in the areas contaminated by the Chernobyl
and Fukushima nuclear disasters. They supplement the knowledge provided in the brief drafted in 2016by IRSN which can be consulted here: What have we learned from the ecological studies carried out
over the long term on the fauna and flora of the territories contaminated by the Chernobyl and
Fukushima accidents?

I - The first sign: the abundance of organisms
To study the populations status, one of the methods frequently used in ecology is the estimation of
the abundance of individuals. A variation in abundance makes it possible to quickly integrate andestimate many deleterious effects of ionizing radiation on populations. Numerous studies have been
carried out on different groups of organisms since the accidents and show contrasting results
depending on the region (Chernobyl, Fukushima) and the group considered (birds, insects, mammals,etc.).
In the Chernobyl region, several research teams have quantified the abundance of large mammals;
however, the results obtained are contradictory (see IRSN 2016, Møller and Mousseau, 2013,
Deryabina et al., 2015, and Gurung et al., 2015) . More recently, old data, collected by teams that had
previously carried out abundance measurements in the field, was re-analysed by integrating a new
estimate of the dose absorbed by organisms (Beaugelin-Seiller et al., 2020). Various parameters that
could influence abundance were taken into account (human activities in the exclusion zone, time ofobservation, and type of forest cover). As the abundance estimation method is based on tracks
observed in the snow, the time elapsed since the last snowfall was also considered as a parameter.
This new study confirms the conclusions of the first analysis carried out in 2013 and shows that themeasured abundances actually depend on the estimated dose (the absorbed dose rates being between
0.0066 and 610 µGy/h): in the most contaminated environments, a 60% decrease in the number ofmammals is observed for an increase of a factor of 10 in the dose (total dose absorbed during thegeneration time of the species considered), this decrease being greater for ‘prey’ (66%) than for‘predators’ (50%).
In Fukushima Prefecture, an a team of American researchers, in cooperation with a Japanese research
centre, also investigated the abundance of mammals and galliform birds following the accident, using
camera traps (Lyons et al., 2020). As in the previous study, the authors analysed the relationships
between radiation levels and the abundances of these organisms and also considered otherparameters, such as human activity (distance to the closest trail and its use), vegetation, altitude, andthe distance to the nearest waterpoint. This is the first study in Fukushima Prefecture to consider the
impact of human activity. This parameter turns out to be very important because the results show that
it is the primary factor affecting the abundance of populations, with a majority of species being moreabundant in areas where human activity is limited. Among other major parameters, the authors also
demonstrated the effect of altitude on population abundance. In contrast, the abundance of wildanimals does not generally seem to depend on the ambient dose. Only one species of pheasant is moreabundant in the most contaminated areas. This study is constrained by the lack of precise
measurements of the dose received by organisms; however, it emphasises the importance ofconsidering the effect of human activities on wild populations to discern the potential effects
induced by exposure to ionizing radiation from those influenced by other factors.
Marine species present on a wide swath of the east coast of Japan, including Fukushima Prefecture,
were monitored in 2012 and 2013. The diversity and abundance of species present in the tidal zone
presented a dependence on the distance to the Fukushima Nuclear Power Plant. Specifically, theabsence of a common mollusc (Thais clavigera) in all of the sites studied near the power plant(Horiguchi et al., 2016) was observed in 2012, while the parts of the coast only affected by the tsunami
were not impacted. More recent monitoring has, on the contrary, revealed an increase in theabundance and diversity of organisms in the areas closest to the plant, four to five years after theaccident (Horiguchi et al., 2020). These studies show the importance of monitoring the abundance of
populations over time, which makes it possible to distinguish the effects resulting from acute exposure(short duration, generally associated with high doses) from those resulting from chronic exposure(longer duration, usually associated with low doses).
The abundance measurement therefore provides a first glimpse of the status of wildlife populations
exposed to ionizing radiation; however, the healthof the individuals observed cannot be assessed using
this measurement alone. In addition, if the reproduction of individuals inhabiting the mostcontaminated areas is impacted, a ‘source-sink’ dynamic falls into place: the weak competition forresources induced due to limited reproduction in the most contaminated areas can promote themigration of individuals towards these areas. This hypothesis has been verified for barn swallows(Hirundo rustica) in the Chernobyl region (the ambient dose rate average on the various sites studied
is between 0.22 and 3.42 µSv/h) (Møller et al., 2006). Populations estimated to be abundant may thus
consist of individuals with different past exposure to ionizing radiation. It is therefore fundamental to
study, more precisely, whether individuals in the most contaminated areas have capacities (survival,
reproduction, etc.) identical to those present in the least contaminated areas.

II - What are the long-term individual effects?
Exposure to ionizing radiation can affect the biological functions of living organisms. Numerous studies
carried out in situ in the Chernobyl region and, to a lesser extent, in the Fukushima region have shownthat radioactive contamination of the environment can generate effects in different species, such asoxidative stress, genetic damage, depression of the immune system, sperm malformations, slowing of
the ability of cells to multiply, partial albinism, and cataracts (Møller and Mousseau, 2006; Geras’kin
et al., 2008; Aliyu et al., 2015; IRSN 2016; Cannon and Kiang, 2020; and, more specifically, e.g. monkey:
Ochiai et al., 2014, Hayama et al. 2017, Urushihara et al. 2018; butterfly: Taira et al., 2014).
However, while these effects are real, in many cases the dose rates described by the authors are often
misjudged. Indeed, in many studies only a portion of the exposure of organisms to radiation is taken
into account: it is limited to external gamma irradiation, and does not consider internal contamination,
the predominant route of exposure for many combinations of organisms and radionuclides (Giraudeau
et al. 2018). It should also be noted that some of the studies carried out in situ did not reveal any
significant negative impacts of ionizing radiation at the individual level (e.g. crustaceans: Fuller et al.,2017, 2018; frog: Giraudeau et al., 2018; fruit fly: Itoh et al., 2018).

These studies confirm that, in addition to the dose absorbed by organisms, the radiosensitivity ofthe species, the biological function studied, the life stage at which the organism is exposed, thetransgenerational effects (Taira et al. 2015), and adaptation capacities must be considered to assess
the effects of ionizing radiation (Shuryak I., 2020).
Recent methodological advances should be highlighted. They involve carrying out a global analysis
without any a priori on the biological functions of organisms, based on changes in gene expression(transcriptomic analysis). This is how Lerebours et al. (2020) studied, thirty years after the Chernobyl
nuclear accident, the effects of radioactive contamination of six Ukrainian and Belarusian lakes on a
species of fish, the perch (Perca fluviatilis); the total dose rate to which the perch were exposed varied
from 0 to 15.7 µGy/h. However, the analysis should be treated with caution, as it was only carried out
on three individuals per lake. However, it highlights disturbances in the expression of genes involved
in biological processes, such as reproduction, the regulation of DNA damage, and lipid metabolism,
consistent with effects on the maturation of oocytes which have been demonstrated in a companion
study on a larger number of fish (Lerebours et al. 2018). Another study without any a priori on the
spleen and liver of bank voles, Myodes glareolus, (Kesäniemi et al. 2019) collected in the Chernobyl
Exclusion Zone and around Kiev, demonstrated effects on the immune system and lipid metabolism. A
dose estimate was performed on each animal. Once again, the importance of these changes on thesurvival of the animals remains difficult to assess but indicates a modification of certain biological
functions linked either to the adaptation of these individuals or to the deterioration of their healthstatus.
However, these global analyses, without any a priori, present a methodological challenge because
the numbers studied are often limited, inter-individual variations can mask the desired effects, andthe databases necessary for data analysis are often non-existent for the wild species studied. The
conclusions that can be drawn at this stage should therefore be treated with caution and crossreferenced with the results and observations obtained by means of approaches deployed at a larger
scale for the study of radiation effects on ecosystems.

III - Do the effects on individuals induce persistent changes within the
population?
Within the same population, not all individuals have the same characteristics (morphology, behaviour,
physiology, etc.). Certain characteristics can be favoured by a selection effect if they allow individuals
to be better able to survive and reproduce in contaminated environments. These individuals are
selected, they have more offsprings and/or a better survival rate. Over the generations, theseoffsprings can come to constitute the majority within the population. Such characteristics are called
adaptations. Since the Chernobyl and Fukushima accidents, several generations of organisms havesucceeded one another and it is probable that such adaptations have taken place. Numerous studies
have thus focused on the adaptation of organisms to ionizing radiation, particularly in the case of theChernobyl accident, and have been compiled in a literature review (Møller et al., 2016). This
adaptation process is often mentioned in the interpretation of certain results obtained in the field
but remains very difficult to demonstrate. To identify an adaptation, it is necessary, in particular, todemonstrate that it persists even after several generations placed in an uncontaminated environment.
This review concluded that, in 2016, only one study could demonstrate this. After irradiating bacteriapresent on swallow feathers from different contaminated sites in the Chernobyl region and from acontrol site in Denmark, the authors of this study (Ruiz-Gonzalez et al., 2016) found evidence of better
bacteria survival and multiplication capacities in those from more contaminated areas (compared tothose from less contaminated areas, ambient dose rates being between 0.03 and 2.9 µGy/h).
Since 2016, new studies seeking to test this adaptation hypothesis have been carried out in the
Chernobyl region. In one of the most recent studies (Arnaise et al., 2020), the authors studied apotential adaptation of fungi infecting the reproductive parts of a flowering plant. By experimentally
irradiating fungi taken from more or less contaminated areas (with ambient dose rates of between
0.03 and 21.03 µSv/h), they demonstrated that fungi from the most contaminated areas are lessresistant to effects of acute and strong experimental irradiation. These results contradict what would
have been expected if the fungi from the most contaminated areas had acquired radiation adaptation
since the accident.
While these adaptations are often seen as beneficial for the viability of wild organisms incontaminated areas, this specialisation can, however, be a handicap in the face of other stresses. In
the event that environmental changes, modifying the living environment of these organisms, appear
in contaminated areas, it is possible that organisms adapted to ionizing radiation are less likely towithstand new environmental changes (for example, those induced by drought and the more frequent
forest fires in the Chernobyl exclusion zone). As such, the destruction of bird nests in the spring by fires
is likely to affect the dynamics of bird populations specifically adapted to organisms in the Chernobyl
exclusion zone (Beresford et al., 2021).
A major direction of research today is therefore to characterise the potential of populations towithstand future changes in the environment.
The adaptation potential of populations is dependent on evolutionary processes involving a persistentmodification of populations over time. This potential can be assessed by measuring genetic diversity.
In the case of the Chernobyl accident, several studies have considered this genetic diversity. A firststudy, carried out on bank voles (Myodes glareolus) (Baker et al, 2017), revealed the existence of agreater genetic diversity in the most contaminated areas (the absorbed dose rates are between 0.29
and 277.92 µGy/h). The authors conclude that there is a greater number of mutations in the mostcontaminated areas but the effect of migration of individuals towards these areas would also make itpossible to obtain such diversity (see ‘source-sink’ dynamics discussed above, Kesäniemi et al., 2018).
On the contrary, recent studies on aquatic crustaceans (Asellus aquaticus) (for absorbed dose rates
between 0.064 and 27.1 µGy/h) and earthworms (for absorbed dose rates between 0.096 and 53µGy/h) do not show a difference in diversity depending on the contamination level (Fuller et al., 2019
and Newbold et al., 2019).
Based on an analysis of the genetic diversity of populations of tree frogs, Eastern tree frogs (Hylaorientalis), a study led by IRSN recently showed, in a similar way to bank voles, an increase in genetic
diversity in the Chernobyl exclusion zone compared to other populations located in less contaminatedareas (absorbed dose rates between 0.007 and 22.4 µGy/h) (Car et al., submitted). By taking into
account the known variations in the genetic diversity of this species in Europe, this study shows that
mutations appear more frequently in the populations in the Chernobyl exclusion zone (an occurrence
frequency approximately one hundred times higher than that known). It is accepted that when a large
number of mutations appear, only a minority allow adaptation to the environment. Indeed, themajority of mutations do not modify the characteristics of individuals or induce deleterious effects.
This high genetic diversity in populations of tree frogs in the Chernobyl exclusion zone has beenmodelled. The models show that this diversity could only be established in populations comprising asmall number of individuals. This study found that among the initial population, only this small numberof individuals, which have the genetic diversity observable today, have adapted to the irradiating
environment. The others have probably disappeared due to the deleterious effects of the mutations
generated by the radiation.
Depending on different characteristics (number of offspring per litter, lifespan, etc.), organisms, suchas large mammals, could be more sensitive than tree frogs to such deleterious mutations. A metaanalysis of studies using various methods and on many groups of organisms was carried out by Møller
and Mousseau (2015) and shows a significant effect of ionizing radiation on the increase in mutations
following the Chernobyl accident. However, they do not show a downward trend in mutations overtime, which could have been expected due to the reduced exposure of the organisms. If this effect ofionizing radiation seems applicable to many groups of organisms, the detailed study of the impact on
genetic diversity concerns, for the moment, only a few of these groups which does not allow us,
today, to have a precise view of the effects of nuclear accidents on the long-term evolution of
populations.

IV - Considering interactions between organisms: what are theconsequences on ecosystem functioning?
The effects of nuclear accidents are generally studied at the scale of an individual or populations of thesame species; however, an ecosystem is not just a juxtaposition of organisms. Indeed, interactions
shape these ecosystems, ranging from the predation of certain mammals to mutualisms between
bacteria and plant roots, including the reproduction of flowering plants through pollinators. A
particularly interesting direction of current research aims to consider these interactions when studying
the effects of ionizing radiation.
One of the first studies on the effects of ionizing radiation on ecosystems and the interactions that
constitute them was carried out by Møller et al. (2012). After observing a decrease in the abundanceof birds and pollinating insects in the most contaminated areas, the authors studied the abundance ofyoung fruit trees. Since these trees are dependent on pollinating insects, which allow the exchange ofpollen to take place, and on birds, which allow the dispersal of seeds, the authors hypothesised thatthese decreases in animal abundance could induce a decrease in young fruit trees. The results of thisstudy actually show a decrease in the abundance of young fruit trees depending on the contaminationof the environment (ambient dose rates are between 0.01 and 379.70 μSv/h) and highlight thedependence of this abundance of young fruit trees to the decrease in pollinating insects and fruiteating birds. Thus, in addition to the potential deleterious effects of ionizing radiation on these fruit
trees, inter-species interactions can induce effects, which are termed ‘indirect effects’.
A particularly interesting and well-documented example of an indirect effect induced by ionizing
radiation is that of pale grass blue butterflies (Zizeeria maha) in Fukushima Prefecture. Since the
Fukushima accident, many studies have focused on this butterfly, demonstrating, for example, anincrease in abnormalities in the wings, eyes, antennae, legs, and mouth parts of the butterflies sampled
in the most contaminated areas (ambient dose rates between 0.08 and 3.09 µSv/h) (Hiyama et al.
2012). To understand the mechanisms behind this effect, researchers reared caterpillars in thelaboratory by feeding them with their host plant, yellow sorrel (Oxalis corniculata). By feedingcaterpillars with yellow sorrel from environments contaminated by the accident, the authors observed
effects (e.g. morphological anomalies) identical to those observed in the field. However, when thecaterpillars are directly contaminated with radionuclides, without going through contamination byyellow sorrel, such effects are no longer observed (Gurung, 2018). The authors of this study therefore
suspected an indirect effect of ionizing radiation linked to yellow sorrel and to the modification of theplant’s composition after contamination. The most recent study on this butterfly-yellow sorrel
interaction indeed shows a change in the nutrient composition of sorrel leaves after contamination,
with, in particular, a decrease in sodium concentration (Sakauchi et al., 2021). The link between themodification of the nutrients present in the leaves and malformations still needs to be studied, but thisexample shows to what extent the very specific effects of contamination by radionuclides on certain
organisms can potentially induce a set of modifications on other organisms, in a cascade, andtherefore on the entire associated ecosystem.
A recent question addressed by Mappes et al. (2019) was to ask if the impact of contamination onorganisms can be modulated by indirect effects. More precisely, can the effects of ionizing radiation
on voles (Myodes glareolus) inhabiting the Chernobyl contaminated zone be modulated by foodsupplementation? Without food supplementation, the authors observed that the abundance of voles
decreased with an increase in exposure to ionizing radiation (ambient dose rates between 0.01 to
95.55 μSv/h). Food supplementation only led to an increase in the abundance of populations inlocations where radioactive contamination was low (<1 μSv/h); in areas with higher contamination,
food supplementation showed no detectable effects. The authors therefore conclude that foodsupplementation can, to a certain extent, mitigate the harmful effects of an environment
contaminated by radionuclides. Once again, this study highlights the importance of considering theinteractions between organisms in the field which can profoundly influence the relationships
between exposure to ionizing radiation and its effects. Since interactions between organisms can be
impacted by exposure to ionizing radiation, this can ultimately affect the functioning of thecontaminated ecosystem. To study these effects, the major functions structuring ecosystem
functioning, such as the decomposition of organic matter (dead plant material, commonly known as'litter'), were studied in the Chernobyl region.
Mousseau et al. (2014) demonstrated a decrease in the rate of leaf litter decomposition and anincrease in the thickness of the litter on the ground related to an increase in ambient radioactivity (theabsorbed dose rate is between 0.09 and 240 μGy/h). These results suggest slower recycling of organic
matter in response to a higher level of ambient radioactivity, possibly due to a negative impact oncommunities of decomposers (bacteria, fungi, invertebrates). Conversely, after conducting the same
type of experiment, a study led by IRSN showed that the leaf litter decomposition rate increased withan increase in the level of radiation absorbed (the absorbed dose rate is between 0.3 and 150 μGy/h)by decomposers (Bonzom et al., 2016). The authors propose two non-exclusive mechanisms to explain
this result: (i) under the effect of moderate stress (here radiological stress) decomposers were able torespond to this disturbance by increasing their survival and reproductive success leading to a greater
abundance of decomposers in radioactively contaminated areas. This phenomenon is known as“hormetic effect” (Calabrese and Baldwin, 2002); (ii) the decomposers communities preferentiallyconsumed the litter from uncontaminated areas rather than the contaminated litter which may havea lower quality that was initially in their habitat (for the purposes of the experiment, theuncontaminated litter was in fact deposited above the contaminated litter in the area chosen for the
study). Certain parameters, such as the contamination gradient, dose estimate, nature of the leaves
used, etc., can explain the contradictory results of these two studies.
The effect of environmental contamination following nuclear accidents on the functioning of
ecosystems is only very marginally considered. However, this is an essential research perspective,
especially due to the fact that the proper functioning of these ecosystems depends on a set of services
these ecosystems provide to human populations (Millennium Ecosystem Assessment, 2005; IPBES,2019). These ‘ecosystem services’ are of primary importance in the more general study of the effects
of global changes and deserves to be more widely integrated into the study of the consequences ofnuclear accidents.

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