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In vivo study: embryotoxicity test
Zebrafish embryos exposed to tested compounds developed lethal and sub-lethal alterations including different abnormalities and unhatching events. LC50 (for mortality rate) and EC50 (for abnormality and unhatching rate) values were extrapolated from concentration–response curves shown in Fig. 1. The rate of dead, abnormal, and/or unhatched specimens was concentration-dependent for all tested compounds (Fig. 1a–c). The lethality of the negative control group was less than 5%. Compounds 4NC and CAT showed the highest toxicity with LC50 values of 8.16 and 10.95 mg/L, respectively, followed by 4,6DNG  > 5NG  > GUA. Experimental LC50/EC50 values and the predicted ones obtained by ECOlogical Structure Activity Relationship (ECOSAR) v2.0 software (https://www.epa.gov/tsca-screening-tools/ecological-structure-activity-relationships-ecosar-predictive-model) based on Quantitative Structure Activity Relationships (QSAR) models showed 4NC and CAT as the most toxic chemicals (Table 2). However, it is important to notice that experimental values for both compounds were approximately two times lower than the predicted ones. This led to the classification of 4NC into the group of molecules toxic to fish (1  GUA).
Figure 3

Recorded sublethal morphological effects in D. rerio embryos/larvae after 48, 72, and 96 h of exposure to CAT, 4NC, GUA, 5NG, and 4,6DNG. Negative control: normally developed embryo at (a) 48, (b) 72, and (c) 96 hpf. During exposure period alterations were manifested as: (d) yolk sac edema (arrow); (e) pericardial edema (asterisk), undeveloped tail region (arrow); (f) hatched fish with malformed spine (arrow); (g) underdeveloped tail and necrosis of its apical part (dashed arrow), rare pigments; (h) pericardial edema (asterisk), scoliosis (arrow), necrosis of the apical part of the tail (dashed arrow), rare pigments, not hatched; (i) scoliosis (arrows), blood accumulation in the brain region (dashed arrow); (j) pericardial edema (asterisk), yolk sac edema (arrow), scoliosis (dashed arrow); (k, l) pericardial edema (asterisk); (m) underdeveloped embryo: underdeveloped head (arrow), tail not detached (asterisk), delay or anomaly in the absorption of the yolk sac; (n) pericardial edema (asterisk), blood accumulation (arrow), not hatched; (o) pericardial edema (asterisk), blood clotting (arrow), not hatched; (p) blood accumulation at the yolk sac (arrow); (r) hatched fish with malformed spine; (s) pericardial edema (black asterisk), blood accumulation above the yolk sac (arrow), swelling of the yolk sac (white asterisk), yolk sac edema (dashed arrow), mild scoliosis. Developmental abnormalities were recorded using LAS EZ 3.2.0 digitizing software (https://www.leica-microsystems.com/products/microscope-software/p/leica-las-ez/).

Full size image

The morphometric measurements (Fig. 4) showed that all tested samples significantly affected sensorial (eye area), skeletal (head height), and physiological (yolk and pericardial sac area) parameters in zebrafish. Significant differences among all treatments with exact p values are presented in Table S2.
Figure 4

Morphometric measurements of D. rerio larvae after 96-h exposure to tested compounds (CAT, 4NC, GUA, 5NG, and 4,6DNG) and control (C). (a) Lateral view showing eye area (EA), head height (HH), yolk sac area (YSA), and pericardial sac area (PSA). Scale bar = 1000 µm. Morphometric parameters are presented by their mean value (b–e; n = 15). The symbol * indicates a significant difference between tested samples and negative control (*p  More

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Vitaly Grishchenko, Yuliia Kulsha, Vasyl Shevchyk & Eugenia Yablonovska-Grishchenko

Smolenskoe Poozerje National Park, Przhevalskoe, Russian Federation
Vladimir Hohryakov, Gennadiy Kosenkov & Ksenia Shalaeva

FSBI Zeya State Nature Reserve, Zeya, Russian Federation
Elena Ignatenko, Klara Pavlova & Sergei Podolski

Polistovsky State Nature Reserve, Pskov, Russian Federation
Svetlana Igosheva & Tatiana Novikova

Ural State Pedagogical University, Yekaterinburg, Russian Federation
Uliya Ivanova, Margarita Kupriyanova, Tamara Nezdoliy, Nataliya Skok & Oksana Yantser

Institute of Mathematical Problems of Biology RAS—the Branch of the Keldysh Institute of Applied Mathematics, Russian Academy of Sciences, Pushchino, Russian Federation
Natalya Ivanova & Maksim Shashkov

Kronotsky Federal Nature Biosphere Reserve, Yelizovo, Russian Federation
Fedor Kazansky & Darya Panicheva

Zhiguli Nature Reserve, P. Bakhilova Polyana, Russian Federation
Darya Kiseleva

Institute for Ecology and Geography, Siberian Federal University, Krasnoyarsk, Russian Federation
Anastasia Knorre

Central Forest State Nature Biosphere Reserve, Tver, Russian Federation
Evgenii Korobov, Elena Shujskaja, Sergei Stepanov & Anatolii Zheltukhin

National Park Bashkirija, Nurgush, Russian Federation
Elvira Kotlugalyamova & Lilija Sultangareeva

State Nature Reserve Kurilsky, Juzhno-Kurilsk, Russian Federation
Evgeny Kozlovsky

Vodlozersky National Park, Karelia, Petrozavodsk, Russian Federation
Elena Kulebyakina & Viktor Mamontov

State Nature Reserve Kivach, Kondopoga, Russian Federation
Anatoliy Kutenkov, Nadezhda Kutenkova, Anatoliy Shcherbakov, Svetlana Skorokhodova, Alexander Sukhov & Marina Yakovleva

South-Ural Federal University, Miass, Russian Federation
Nadezhda Kuyantseva

Saint-Petersburg State Forest Technical University, St. Petersburg, Russian Federation
Pavel Lebedev

Astrakhan Biosphere Reserve, Astrakhan, Russian Federation
Kirill Litvinov

FSBI United Administration of the Lazovsky State Reserve and National Park Zov Tigra, Lazo, Russian Federation
Lidiya Makovkina, Aleksandr Myslenkov & Inna Voloshina

State Nature Reserve Tungusskiy, Krasnoyarsk, Russian Federation
Artur Meydus, Julia Raiskaya & Vladimir Sopin

Krasnoyarsk State Pedagogical University named after V.P. Astafyev, Krasnoyarsk, Russian Federation
Artur Meydus

Institute of Geography, Russian Academy of Sciences, Moscow, Russian Federation
Aleksandr Minin

Koltzov Institute of Developmental Biology, Russian Academy of Sciences, Moscow, Russian Federation
Aleksandr Minin

Carpathian National Nature Park, Yaremche, Ukraine
Mykhailo Motruk

State Environmental Institution National Park Braslav lakes, Braslav, Belarus
Nina Nasonova

National Park Synevyr, Synevyr-Ostriki, Ukraine
Tatyana Niroda, Ivan Putrashyk, Yurij Tyukh & Yurij Yarema

Pasvik State Nature Reserve, Nikel, Russian Federation
Natalja Polikarpova

Mari Chodra National Park, Krasnogorsky, Russian Federation
Tatiana Polyanskaya

State Nature Reserve Vishersky, Krasnovishersk, Russian Federation
Irina Prokosheva

State Nature Reserve Olekminsky, Olekminsk, Russian Federation
Yuri Rozhkov, Olga Rozhkova & Dmitry Tirski

Crimea Nature Reserve, Alushta, Republic of Crimea
Marina Rudenko

Forest Research Institute Karelian Research Centre, Russian Academy of Sciences, Petrozavodsk, Russian Federation
Sergei Sazonov, Lidia Vetchinnikova & Juri Kurhinen

Black Sea Biosphere Reserve, Hola Prystan’, Ukraine
Zoya Selyunina

Institute of Physicochemical and Biological Problems in Soil Sciences, Russian Academy of Sciences, Pushchino, Russian Federation
Maksim Shashkov

State Nature Reserve Nurgush, Kirov, Russian Federation
Sergej Shubin & Ludmila Tselishcheva

Caucasian State Biosphere Reserve of the Ministry of Natural Resources, Maykop, Russian Federation
Yurii Spasovski

National Nature Park Vyzhnytskiy, Berehomet, Ukraine
Vitalіy Stratiy

National Park Khvalynsky, Khvalynsk, Russian Federation
Guzalya Suleymanova

State Research Center Arctic and Antarctic Research Institute, Saint Petersburg, Russian Federation
Aleksey Tomilin

Information-Analytical Centre for Protected Areas, Moscow, Russian Federation
Aleksey Tomilin

State Nature Reserve Malaya Sosva, Sovetskiy, Russian Federation
Aleksander Vasin & Aleksandra Vasina

Krasnoyarsk State Medical University named after Prof. V.F.Voino-Yasenetsky, Krasnoyarsk, Russian Federation
Vladislav Vinogradov

Surhanskiy State Nature Reserve, Sherabad, Uzbekistan
Tura Xoliqov

Mordovia State Nature Reserve, Pushta, Russian Federation
Andrey Zahvatov

Centre for Biodiversity Dynamics, Department of Biology, Norwegian University of Science and Technology, Trondheim, Norway
Otso Ovaskainen

The data were collected by the 195 authors starting from M.A. and ending with T.Z. in the author list. J.K., E.M., C.L., G.T. and E.G. contributed to the establishment and coordination of the collaborative network and to the compilation and curation of the resulting dataset. T.R., O.O., L.A., M.H. and M.d.M.D. conceived the idea behind the current study and wrote the first draft of the paper, with O.O. conducting the analyses. All authors provided useful comments on earlier drafts. More