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<I>Anethum Graveolens</I>

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<I>Anethum Graveolens</I> Bukharov, Baleev, Soldatenko, Musaev, Kezimana and Priyatkin (2021). Seed Science and Technology, 49, 1, 7-17. https://doi.org/10.15258/sst.2021.49.1.02 Impacts of high temperature on embryonic growth and seed germination of dill (Anethum graveolens) Alexander F. Bukharov1, Dmitry N. Baleev2, Aleksey V. Soldatenko3, Farhad B. Musaev3, Parfait Kezimana4 and Nikolay S. Priyatkin5* 1 All-Russian Scientific Research Institute of Vegetable Growing, Branch of the «Federal Scientific Vegetable Center», Moscow Region, 140153, Russia 2 All-Russian Research Institute of Medicinal and Aromatic Plants, 7, Grina street, Moscow, 117216, Russia 3 Federal Scientific Vegetable Center, 14, Selektsionnaya Street, Odintsovo District, Moscow Region, 143080, Russia 4 Peoples Friendship University of Russia (RUDN University), 6, Miklukho-Maklaya Street, Moscow, 117198, Russia 5 Agrophysical Research Institute, 14, Grazhdanskiy pr., St. Petersburg, 195220, Russia * Author for correspondence (E-mail: [email protected]) (Submitted July 2020; Accepted December 2020; Published online January 2021) Abstract Temperature is the main exogenous factor that determines the growth, development and productivity of agricultural plants, including seed germination. In this work, we studied the thermosensitivity of heterogeneous dill seeds (Anethum graveolens). The study aimed to study the short-term (1-5 days) effect of high temperature (40°C) on the growth of the embryo and germination of dill seeds formed on different umbels of the mother plant. The growth of the embryo had a significant effect on seed germination (r = 0.976; P < 0.001). Seeds collected from primary umbels have a more developed embryo and can overcome the effects of thermal stress. We also found that the duration of heat exposure at 40°C, leading to a decrease in viability, can vary from 1-5 days for seeds from primary umbels, to 1-2 days for seeds from secondary umbels, which, in our opinion, is due to the heteromorphism. The influence of the place of seed formation on the mother plant and the high temperature during germination is critical for embryo growth and dill seeds’ germination. Keywords: Anethum graveolens L., embryo growth, germination, seed heterogeneity, thermal sensitivity Introduction One of the unfavourable abiotic factors, high temperature, affects plant growth and development at different stages of development, including during seed germination. Temperature is one of the main factors determining the productivity of many cultivated plants (Maraghni et al., 2010; Wen, 2015; Chitwood et al., 2016), including those in the Apiaceae family (Nascimento et al., 2013a, b). An increase of 10-15°С above the optimum causes stress, manifested in changes in metabolic reactions and physiological processes (Wang et al., 2004; Tariq et al., 2010). © 2021 Bukharov et al. This is an open access article distributed in accordance with the Creative Commons Attribution Non Commercial (CC BY-NC 4.0) license, which permits others to distribute, remix, adapt, build upon this work non-commercially, and license their derivative works on different terms, provided the original work is properly cited and the use is non-commercial. See: https://creativecommons.org/licenses/by-nc/4.0 7 A.F. BUKHAROV, D.N. BALEEV, A.V. SOLDATENKO, F.B. MUSAEV, P. KEZIMANA AND N.S. PRIYATKIN Among the crops of the Apiaceae family, seed heterogeneity due to seed position on the mother plant is widespread (Gray and Steckel, 1985; Bianco et al., 1994). Seed heterogeneity, when a single plant produces seeds that differ in size, weight, colour, mor- pho logy, anatomy, germination and/or other characteristics (Sun et al., 2009; Panayotov, 2009; Yao et al., 2010; Cao et al., 2015), is widely represented in the plant world and is inherent in both wild and cultivated forms. In many plant species, significant variations in seed size have been detected (Moles and Westoby, 2006; Pinto et al., 2018), some characterised by discrete variability (Venable et al., 1987), while others show a continuous variation in seed traits (Stanton, 1984; Agren, 1989). Many authors suggest that seed heterogeneity is a consequence of crop adaptation to environmental factors (Galloway et al., 2009; Weber, 2009; Dyer et al., 2010), such as salinity, high soil moisture and stressful air temperatures (Berger, 1985; Zhang, 1995). There are several types of seed heterogeneity: genetic, due to the combination of hereditarily unequal parental gametes (Smith and Fitzsimmons, 1965); maternal, due to the difference in the seeds’ position on the mother plant, which results in different developmental conditions (Wolf and Wade, 2009); and ecological, due to the interaction of developing seeds with environmental conditions (Wardlaw et al., 1989). Earlier studies have shown that seed size and weight in Pastinaca sativa L. decrease with increasing umbel order (Hendrix, 1984), while the variation in seed size within an individual plant of Lomatium grayi was 16% (Thompson, 1984). Several researchers have studied the effect of seed heteromorphism on their quality in Apiaceae plants, and they have shown that the quality (Corbineau et al., 1995; Panayotov, 2010) and dormancy onset (Thomas et al., 1978) of seeds change depending on the seed position on the mother plant. Dill (Anethum graveolens L.) is an annual herbaceous vegetable and aromatic plant of the Apiaceae family, which has been used for medical and culinary purposes since ancient times (Radulescu et al., 2010). Wild dill can be found in South-West and Central Asia, and in South-East Europe, while the cultivated crop is well-known and used all over the world (Bailer et al., 2001). Dill forms a single, straight, branched stem up to 1.5 m in height, with thin alternating cauline leaves. Flowers are collected in small umbels with a diameter of 20-90 mm, which are grouped into a large umbel that is up to 150-200 mm in diameter. In the Moscow region, the flowering of dill plants is usually in June-July, and fruits (cremocarp) ripen in July-September. Dill seeds are ovoid or broadly oval, about 3-5 mm long and 1.5-3.5 mm wide (Warrier et al., 1994). Dill is one of the most important spice vegetable crops grown throughout Russia and improving seed quality is undoubtedly relevant to increase production. A new late-ripening and fruticose variety ‘Kentavr’ can form ripe seeds in primary and secondary umbels in the Moscow region conditions. It is known that the position of the seed on the plant or even within the umbel (position effect) can affect its morphological structure, weight, size, chemical composition, germination and dormancy (Matilla et al., 2005). Previously, we conducted a study on seed development depending on the position in the umbels and on the mother plant of this variety and identified seed heterogeneity in the length of the endosperm and embryo (Bukharov et al., 2017). With the increasing frequency of abnormal weather conditions, it is relevant to study the adaptive responses of heterogeneous seeds to stress factors. This study aimed to investigate the differences in the thermosensitivity of heterogeneous dill seeds. 8 IMPACTS OF HIGH TEMPERATURE ON DILL EMBRYONIC GROWTH We assumed that seeds from primary and secondary umbels react differently to high temperatures during germination, as seeds with a more developed embryo can overcome this stress factor. The effect of high temperature has been studied on carrots (Pereira et al., 2008; Nascimento et al., 2012), but such studies on dill crops are rare (Holubowicz and Morozowska, 2011). Thus, the main goal of our research was to find out how the short-term (1-5 days) effect of high temperature (40°C) affects embryo growth during germination and the germination of heterogeneous dill seeds. The specific objectives of this study were to (a) describe the phenology of embryo growth and germination; (b) characterise the germination of seeds formed in various umbels on the mother plant in response to the effect of a critical temperature (40°C); and (c) determine the time of exposure to the critical temperature for embryo growth and germination of heterogeneous dill seeds. Materials and methods Studies were conducted in 2015 and 2016 at the All-Russian Scientific Research Institute of Vegetable Growing – Branch of the “Federal Scientific Vegetable Center” (Moscow Region, N55°36'E38°1'). Dill seeds of the cultivar ‘Kentavr’ were collected from primary or secondary umbels. At the flowering phase, the plant is 1.0-1.1 m in height, sprawling with many leaves, and its umbel is large, convex and multi-radiate. Seeds were collected in an open field in 2015 and 2016, and harvested 50 days after the flowering of the primary umbels. Umbels were cut from 30 plants for each variant and dried indoors (22- 25°C) for 3-5 days. The dried seeds were placed in paper envelopes and stored in a sealed plastic container with silica gel at 4-5°C for six months. Separate studies were carried out for the seeds from each year. Incubation of seeds under high-temperature conditions For the analysis of the impact of short-term high-temperature stress, imbibed seeds from primary and secondary umbels were incubated at 40°C for 0, 1, 2, 3, 4 or 5 days. For the embryo growth analysis, seeds (four replicates of 300 seeds each) were placed in Petri dishes (diameter 100 mm, height 20 mm) on two sheets of filter paper moistened with distilled water and sealed with parafilm. Each of the 40 Petri dishes used was placed in a thermostat TC 1/80 (SKTB SPU, Russia) with controlled temperature and light. In parallel, seeds were incubated for the analysis of the effect of high temperature on seed germination, as described above. After incubation, seeds were transferred to standard conditions (t = 20°C) to study embryo growth and seed germination. These conditions were chosen as they are optimal for the germination of seeds of many species (Nikolaeva, 1969).
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