110 Annales Universitatis Paedagogicae Cracoviensis Studia Naturae, 5: 110–128, 2020, ISSN 2543-8832 DOI: 10.24917/25438832.5.8 Katarzyna Lipniak1, Angelika Kliszcz2* 1Cracow Institute of Development and Education, Wielicka 42/105 St., 30-552 Kraków, Poland 2University of Agriculture in Krakow, Department of Agroecology and Plant Production, Mickiewicz 21 Ave, 30-120 Kraków, Poland; *angelika.kliszcz@student.urk.edu.pl Allelopathic effect of goosefoot on germination and early stage growth of triticale and radish Introduction Weeds are plants that are very well adapted to growth and development in changing environmental conditions. �eir proliferation is a  signi�cant and growing problem for agriculture. �ey have the ability to produce an ample amount of seeds which are characterised by prominent vitality and easy germination. �ese plants demonstrate a wide range of temperature tolerance and variable soil conditions. �ey use nu- trients in larger quantities than crop plants and they can appear in secondary weed infestation. One example is goosefoot (Chenopodium album L.). It is an annual plant that grows to about one meter in height (Paczyńska, 2016). In various classi�cation systems it belongs to the goosefoot family (Chenopodiaceae Vent.) or amaranthus (Amaranthaceae Juss.). C. album has been cultivated in Europe since ancient times. In Poland, it occurs in the lowlands as well as in lower mountain regions, and it o�en grows on the fallow, �elds or gardens (Sudnik-Wójcikowska, 2011). Goosefoot has a  pile root and is branched. Its stem also branches strongly and sometimes has red discoloration. �e whole plant is pubescent (Czubiński, Paradowski, 2014). �e leaves take on a diamond shape or are lanceolate, with a serrated edge and a wedge-shaped base. C. album begins �owering in July, which can last until November. Its �owers are small and pale green. A�er �owering, it produces fruit in the form of a small nut with black seeds. Goosefoot seeds can easily adapt to all weather conditions, even retain- ing their ability to germinate for several years (Klaaβen, Freitag, 2004). �is plant is wind-proof and reproduces and develops very quickly, thus taking valuable nutrients and water from crop plants (Domagała-Świątkiewicz, 2007). C. album contains vi- tamins A, B1, B2 and C and microelements. It is also rich in �avonoids, essential A llelopathic effect of goosefoot on germ ination and early stage grow th of triticale and radish 111 oils, carbohydrates and proteins (Byłka, Kowalewski, 1997; Dutt et al., 2003; Jardim et al., 2008; Usman et al., 2010; Gęsiński, Nowak, 2011). Its seeds are abundant in valuable fats and albumins. Shoots, older leaves and seeds contain oxalic acid, sapo- nins, phenols, lignans and alkaloids (Cutillo et al., 2004, 2006; Lavaud et al., 2007; Laghari et al., 2011) Crop plants and weeds are a community with an additive system of components in which individuals compete for limited resources of the habitat, especially for water, nutrients and light. Competition results in the adjustment and weakening of compet- ing organisms, which manifests as a  reduction in the amount of biomass produced, in the size of individual organs and in seed yield. In cases of strong competitive impacts, individuals may even die and be eliminated from the community (Begon et al., 1999). In the literature, the high allelopathic potential of C. album is widely dis- cussed (Batish et al., 2006; Jafari, Kholdebarin, 2002; Laghari et al., 2011). Although, Reinhardt et al. (1994) pointed out, this plant did not exert an inhibitory e�ect on radish germination when milled dry shoot biomass was added to the soil (1% w/w) or when radish seeds were treated with aqueous soil extracts from pots that previously contained mature C. album plants. In natural conditions, the allelopathic actions of C. album are not only limited to plant-plant interactions. C. album plants strongly prop- agate soil microorganism growth (like Anabena genus, a nitrogen-�xing cyanobacte- ria) and their aqueous extracts can enhance the growth of Bradyrhizobium japonicum Kirchner (Vokou et al., 2006). �e aim of the study was to examine the e�ect of aqueous extracts from dry goosefoot (Chenopodium album L.) plants on germination and early stage growth of triticale grains (×Triticosecale Wittm. ex A.Camus) and radish seeds (Raphanus sa- tivus L.). �e e�ect of di�erent percent concentrations of C. album extracts on (1) germination index values, (2) percent inhibition of seedling growth, (3) fresh and dry seedling mass, relative water content and percent water content in 7-day triticale and radish seedlings and (4) the degree of cell membrane destabilisation by measuring the electrolyte leakage were determined. Material and methods Plant material Triticale (×Triticosecale) grains and radish (Raphanus sativus) seeds were used in the experiment. Goosefoot (Chenopodium album) shoots in the vegetative phase were collected in spring 2019, from the southern Poland stand (49°59ʹ10ʺN 20°03ʹ42ʺE) and dried in laboratory conditions. Plant material was stored in the dark to avoid photochemical destruction of allelopathic compounds. K at ar zy na L ip ni ak , A ng el ik a K lis zc z 112 Preparation of extracts Dry material (goosefoot shoots) was ground in a mortar and distilled water was add- ed, in the following amounts: (I) 0.5% extract: 0.5 g dry material + 95.5 ml distilled water, (II) 1.0% extract: 1 g dry material + 99 ml distilled water, (III) 1.5% extract: 1.5 g dry material + 98.5 ml distilled water and (IV) 2.0% extract: 2 g dry material + 98 ml distilled water. �e preparations were le� for 24 hours in the dark at approx- imately 25°C to allow for extraction of the compounds. A�er one day, the extracts from dry C. album plants were strained and stored in a refrigerator at 8°C ± 2°C for the duration of the experiment. Seed germination 25 pieces of triticale grains and radish seeds (a�er washing with running water for 30 minutes, and then 3 times with distilled water) were placed with a sterile tweezer on Petri dishes (Ø 9 cm) covered with a triple layer of �lter paper, moistened with 5 ml of the appropriate aqueous extract of C. album plants in the concentrations: 0.5%, 1.0%, 1.5% or 2.0%. For the control, distilled water was used. During the experiment, all grains and seeds were placed in the dark, at room temperature. Every 24 h for 7 days the number of germinated grains and seeds was counted. Grains and seeds were consid- ered germinated when their germinal root was equal to half the size of a grain or seed. Germination parameters A�er 7 days of the experiment, the e�ect of C. album aqueous extracts on the germi- nation capacity of triticale grains and radish seeds was evaluated. A germination index – GI (AOSA, 1983), speed of emergence – SE, seedling vigour index – SVI (Islam et al., 2009), coe�cient of the rate of germination – CRG (Chiapusio et al., 1997) and time required for 50% germination – T50 (Farooq et al., 2006) were assessed. Biometric analysis Triticale and radish seedling length was measured using a calliper with an accuracy of 1 mm. �e e�ect of goosefoot extracts on seedling growth was determined according to Islam and Kato-Noguchi (2012). Fresh and dry mass, relative water content and percentage water content Fresh mass (FM) of triticale and radish seedlings was determined with a balance (Ohaus Adventurer Pro, USA). To obtain the dry mass (DM), the plant material was dried for 48 h at 105°C in a dryer (WAMED SUP 100, Poland) and then weighed. �e relative water content (RWC) and the percentage content were determined based on the mass values. A llelopathic effect of goosefoot on germ ination and early stage grow th of triticale and radish 113 �e RWC was determined according to the method described by Mullan and Pi- etragalla (2012). Brie�y, an individual seedling was weighed for FM and incubated for 24 h at 25°C in vials with 10 ml distilled water for saturation of plant tissues with wa- ter. A�er the incubation time, the turgor seedling mass (TM) was determined. �en, each was dried at 105°C for 48 h in the laboratory oven (WAMED SUP 100, Poland) and DM weighed. �e RWC parameter for every seedling was calculated according to the formula: RWC = [(FM − DM) / (TM − DM)] × 100. �e percentage of water content (% H2O) was determined based on the mass values according to the formula % H2O = 100 − [(DM × 100) / FM]. Electrolyte leakage Cell membrane permeability was measured by electrolytze leakage in triticale and rad- ish seedlings according to the method used by Możdżeń et al. (2018). Statistical analysis Experimental results were compiled in Microso� Excel. Additionally, statistical anal- ysis was performed using one-way ANOVA/MANOVA. To assess the signi�cance of di�erences between the means ± SD (n = 3), Duncan’s test at p ≤ 0.05 was used. �e data was analysed with the STATISTICA program (StatSo�, Inc. 2018, Data Analysis So�ware System, version 13.1). Results Germination indexes �e germination capacity of triticale grains (×Triticosecale) from the control group (distilled water) a�er 2 days was 100%. A similar result was obtained for seeds germi- nating on the aqueous extracts of dry Chenopodium album at a concentration of 0.5%, where the germination capacity a�er 3 days was 90%. Higher concentrations of goose- foot extracts (1.0%, 1.5% and 2.0%) inhibited the germination of grains. Regardless of the concentration of extracts, the largest number of newly germinated triticale grains was observed on the third day of the experiment. Similar results were observed for radish (Raphanus sativus) seeds. �e highest percentage of germinated seeds was recorded a�er 4 days for the control sample and a�er 6 days for the 0.5% aqueous ex- tracts. With increasing concentrations, a clear reduction in the number of germinated seeds was observed. Radish seeds watered with 2.0% extracts exhibited the lowest ger- mination capacity (Tab. 1 – Appendix 1). �e coe�cient of rate of germination (CRG) index for triticale grains slightly de- creased compared to control with increasing aqueous goosefoot extract concentra- tions. �e only statistically signi�cant di�erences observed for this index, for grains, K at ar zy na L ip ni ak , A ng el ik a K lis zc z 114 were between the control and 2.0% extracts. Compared to the control, the CRG for radish seeds clearly decreased in each of the C. album extracts used. A  signi�- cant decrease in the CRG index was demonstrated even at a  concentration of 1.0% (Tab. 2 – Appendix 1). �e shortest time needed to reach 50% germinated seeds (T50) for triticale and radish seeds was in the control case. For aqueous C. album extracts, the T50 values for each of the extracts increased with the concentration of allelopathic compounds (Tab. 2 – Appendix 1). �e germination index (GI) reached higher values for rad- ish seeds compared to triticale grains. Regardless of the type of seeds studied, the GI values decreased with increasing concentrations of aqueous C. album extracts. Com- pared to the control, the lowest GI was observed for grains and seeds germinating in Petri dishes saturated with 2.0% goosefoot extracts. �e speed of emergence (SE), regardless of the type of studied seeds, reached its highest values in the control sample (Tab. 2 – Appendix 1). For C. album extracts, the SE index values decreased for both triticale and radish seeds with an increase in the concentration of extracts. For radish seeds treated with extracts (1.5% and 2.0% con- centrations) a clear inhibition of germination was observed. Compared to the control, the seed vigour index (SVI), irrespective of the organ and type of seeds, signi�cantly decreased with increasing concentration of C. album extracts (Tab. 3 – Appendix 1). �e lowest values were observed in seedlings watered with 2.0% extracts. Biometric analysis Biometric analysis of whole triticale and radish seedlings revealed signi�cant inhibi- tion of growth in the presence of aqueous extracts from C. album plants (Tab. 4 – Ap- pendix 1; Fig. 1–2 – Appendix 2). Stimulation of seedling growth in relation to con- trol was observed only for radish seedlings watered with 0.5% goosefoot extract. �e root length of triticale and radish seedlings was similar in all studied samples. With increasing concentration of allelopathic compounds in aqueous extracts, inhibition of root growth was observed. �e exception was for R. sativus seedlings watered with 0.5% C. album extract, which resulted in a stimulatory e�ect on the root length of seed- lings. �e inhibition percentage index of growth (IP), expressed as a percentage of the control value, reached higher positive values for the triticale and radish above-ground organs with increasing concentration of goosefoot extract (Tab. 4 – Appendix 1). �e only exception were radish seedlings watered with 0.5% extract; compared to controls, they reached negative values, which indicated stimulation of hypocotyl growth. A llelopathic effect of goosefoot on germ ination and early stage grow th of triticale and radish 115 Fresh and dry mass, relative water content and percentage water content �e aqueous extracts from C. album plants had an inhibitory e�ect on the FM values of triticale and radish seedlings. Compared to the control sample, when the concentra- tion of the extracts increased, a signi�cant decrease in the mass values was observed. �e lowest values for this parameter were found in seedlings watered with 2.0% ex- tract. �e exception was radish seedlings treated with 0.5% extract, which showed a signi�cant increase in FM, compared to control seedlings (Tab. 5). �e DM of triticale seedlings reached di�erent values depending on the concen- tration of the extract. Compared to the control, the highest values were found in seed- lings watered with 0.5% extract and the lowest for seedlings grown in Petri dishes with 1.0% and 2.0% goosefoot extract. In the case of radish seedlings, DM increased for each extract concentration, in relation to the control (Tab. 5). Tab. 5. Fresh and dry masses, percentage water content and relative water content of triticale (×Triti- cosecale Wittm. ex A.Camus) – (A) and radish (Raphanus sativus L.) – (B) seedlings watered with the aqueous extracts of Chenopodium album L. shoots in di�erent concentrations (0.5%, 1.0%, 1.5%, 2.0%) and control conditions Pa ra m et er Concentration of aqueous extracts [%] Control 0.5 1.0 1.5 2.0 A B A B A B A B A B FM 0.2531a±0.03 0.0988b ±0.02 0.2196ab ±0.07 0.1587a ±0.01 0.1813b ±0.03 0.0673c ±0.03 0.1699c ±0.02 0.0283d ±0.02 0.1018d ±0.01 0.0197e ±0.005 DM 0.0251c±0.00 0.0028d ±0.001 0.0796a ±0.06 0.0050a ±0.00 0.0227c ±0.01 0.0043b ±0.00 0.0577b ±0.08 0.0025d ±0.00 0.0267c ±0.01 0.0039c ±0.002 WC 90.09a±0.29 97.10a ±0.47 57.88de ±39.07 96.83a ±0.66 87.52ab ±1.51 92.80a ±2.65 68.20d ±39.01 90.82a ±2.61 73.80c ±4.67 78.05b ±17.19 RWC 79.61a±8.89 58.50b ±15.00 60.73bc ±29.84 70.51a ±3.61 66.74b ±6.23 57.64b ±12.44 50.54c ±29.93 75.28a ±53.29 60.82bc ±14.87 51.77b ±35.39 mean values ±SD (n = 3) in row marked with letters a, b, c di�er signi�cantly according to Duncan’s test at p ≤ 0.05; FM – fresh mass [g], DM – dry mass [g], WC – water content [%], RWC – relative water content [%] �e total water content of triticale seedlings decreased signi�cantly when extract concentration increased, compared to the control values (Tab. 5). In the case of rad- ish seedlings, the values for this parameter changed slightly. In seedlings watered with 2.0% extract, a  signi�cant decrease in the water content was found. �e RWC in the tested triticale seedlings decreased with the increasing extract concentration, compared to the control. For radish seedlings, the values for this parameter di�ered depending on the interaction with allelopathic compounds concentrated in aqueous extracts from C. album. K at ar zy na L ip ni ak , A ng el ik a K lis zc z 116 Electrolyte leakage �e electrolyte leakage measurements revealed an increase in cell membrane desta- bilisation under the in�uence of goosefoot extracts (Fig. 3). Regardless of the type of seedling, a statistically signi�cant increase in electrolyte leakage occurred along with increasing extract concentration. �e highest degree of cell membrane disorganisa- tion was found for triticale and radish seedlings treated with 2.0% C. album extract. Fig. 3. Electrolyte leakage from cell membranes of triticale (×Triticosecale Wittm. ex A.Camus) and rad- ish (Raphanus sativus L.) seedlings watered with the aqueous extracts of Chenopodium album L. shoots in concentrations 0.5%, 1.0%, 1.5%, 2.0% and control conditions; mean values ±SD (n = 3) marked with letters a, b, c di�er signi�cantly according to Duncan’s test at p ≤ 0.05 Discussion Seed germination is a complex process that includes both catabolic and anabolic re- actions and biochemical transformations. It consists of processes occurring inside the seed that lead to the activation of the embryo (Nonogaki et al., 2010). �e germination is controlled by external (environmental) and internal (genetic and hormonal) fac- tors realised as a function of time. �e exogenous factors include water, temperature and light (Benech-Arnold et al., 2000). �e endogenous factors are hormones, growth and development regulators, and reactive oxygen species content. In addition, various chemical substances that occur in nature, e.g. nitrogen oxides, butenolide derivatives (Janas et al., 2010), glyceronitrile (Downes et al., 2013) and allelopathic compounds secreted by neighbouring plants (Zandi et al., 2018, 2019; Puła et al., 2020) have im- pact on germination. A llelopathic effect of goosefoot on germ ination and early stage grow th of triticale and radish 117 �is research carried out on C. album con�rms that, during germination, grains and seeds become sensitive to allelopathic compounds (Vokou et al., 2006; Valizadeh, Mirshekari, 2011; Konieczna et al., 2018). �e germination capacity of triticale grains and radish seeds from the control sample was identical to the sample treated with 0.5% C. album aqueous extract. Meanwhile, in other shoot extracts it turned out that the higher the concentration of allelopathic substances the smaller the number of ger- minated grains and seeds (Tab. 1–2 – Appendix 1). For GI values, a similar result was observed. It reached higher values for radish seeds, compared to triticale grains. In the control groups, the highest values of SE index were observed. �e lowest SE values were observed for grains and seeds treated with allelochemical compounds that had accumulated in the 1.5% and 2.0% extracts (Tab. 2 – Appendix 1). For CRG, values clearly decreased with increasing concentration of aqueous C. album extract, compared to the control sample (Tab. 2 – Appendix 1). �e SVI reached its lowest values for seedlings watered with 2.0% shoot extract, regardless of the type of organ and seeds (Tab. 3 – Appendix 1). Similar observations in terms of inhibition of seed germination in the presence of aqueous extracts from C. album shoot for Ra- phanus sativus L. were observed by e.g. Mallik et al. (1994) and Konieczna et al. (2019). Di�erent degrees of seed sensitivity to allelopathic compounds may result from the size of diaspores and the thickness of seed coats (Sołtys et al., 2012). According to Vaughn and Spencer (1993), large-sized seeds show higher stress tolerance. Możdżeń and Rzepka (2016) and Mazur (2019) con�rmed the important protective role of seed coat in the germination and early stages of growth of Vicia faba L. and Phaseolus vul- garis L. �e mechanism of action of allelopathic compounds on seed germination is also associated with disturbances in the phytohormonal balance, slowing down the activation of spare materials, as well as the induction of oxidative stress (Krasuska et al., 2014). Biometric analysis of underground and above-ground organs of triticale and rad- ish seedlings revealed that aqueous extracts from C. album shoots inhibited their growth, as the concentration of allelophatins in the extracts increased (Tab. 4 – Ap- pendix 1). �e exception turned out to be radish seedlings watered with 0.5% extracts. �is extract stimulated their growth, compared to seedlings from the control. Most likely contained in the leaves and stems of C. album, nitrates, among others phenols, saponins and alkaloids, negatively a�ected the germination and early stages of growth of triticale and radish seeds (Cutillo et al., 2004, 2006; Lavaud et al., 2007; Czubiński, Paradowski, 2014). Allelopathic substances inhibit cell division and cell lengthening by limiting pro- ton transport from the cytoplasm to the apoplast (Burgos et al., 2004). �ey reduce the uptake of micro and macroelements by changing the hydraulic conductivity of cell membranes. One of the �rst e�ects of allelophatic compounds at a cellular level is to K at ar zy na L ip ni ak , A ng el ik a K lis zc z 118 reduce the transmembrane electrochemical potential of cell membranes. Membrane depolarisation causes disturbances in the transport of anions and cations, which is as- sociated with increased permeability of these structures. Under stress conditions, (1) inhibition of phosphorus, potassium, magnesium and nitrate ion intake, (2) modi�ca- tion of membrane proteins, (3) lipid oxidation by the presence of free radicals due to the reduction of catalase and peroxidase activity and (4) disorders in the functioning of channels, membrane conveyors and proton pumps occur. Damage to the cell mem- branes depends on, among other things, the concentration and solubility of allelo- pathic substances and the pH of the environment (Einhellig, 2004). In the experiment, di�erences in the electrolyte leakages from triticale and radish seedlings proved that allelopathic compounds released from goosefoot shoots clearly increased the degree of cell membrane destabilisation (Fig. 3). Studies on the allelopathic properties of C. album showed that the presence of goosefoot reduced the quantity and quality of various crop plant species (Krop� et al., 1992; Salam et al., 2014). For example, in Zea mays L., Glycine max (L.) Merr., Solanum L. section Lycopersicon (Mill.) Wettst., Avena L., Hordeum L., Medicago L., and Beta vulgaris L. subsp. vulgaris, at a density of 172 to 300 plants per m2, goosefoot caused crop losses from 6 to 58% (Staniforth, Lovely, 1964; Sibuga, Bandeen, 1980; Shurtle�, Coble, 1985; Torner et al., 1995; Ngouajio et al., 1999). Dyck and Liebman (1995) attributed uncontrolled C. album populations to as much as 59% reduction in yields. In tomato cultivation, goosefoot caused a  36% reduction in the quality of marketable fruit (Bhowmik, Reddy, 1988). For barley yield, changes from 23 to 36% were attributed to its competition with C. album plants (Conn, �omas, 1987). C. album caused an approximate 60% reduction in grain yield during the oat growing season and a 23% reduction in lucerne biomass yield (Lapointe et al., 1985). �is study also con�rmed a negative e�ect of C. album extract on the masses and water content of triticale and radish seedlings (Tab. 5). A decrease in mass accumulation was observed with increasing extract concentration, regardless of the type of seeds. Only for rad- ish seedlings was an increase in the DM value for each of the extract concentrations found. �e intense and negative allelopathic e�ect of goosefoot extracts on triticale grains and radish seeds most likely resulted from the type of extracts used in the experiment. Leaves are organs that accumulate the largest amount of allelopathic compounds, and roots contain the lowest concentration (Kryzeviciene, Paplauskiene, 2004). Goosefoot is a common weed in Poland. In limiting C. album, regular care of ar- able land in connection with simultaneous prevention of weed control is important. Otherwise, C. album plants le� without interference quickly cover large areas (even eliminating crop species), will use the nutrients intended for the crop plants and sharply increase their seed bank in the soil. �is problem especially a�ects plants cul- A llelopathic effect of goosefoot on germ ination and early stage grow th of triticale and radish 119 tivated in wide rows, such as maize, sugar beets and potatoes. �e weed economic thresholds determine the number of weed plants per unit area at which the crop yield loss will be greater than the total costs of the plant protection treatments. For maize, the threshold is 2 pcs of C. album per m2 and for sugar beet 5 plants per 30 m in a row (�e threshold of pests…, 2020). In cereal crops (both spring and winter forms), besides the negative potential from released allelochemicals and competition for nutrients, the timing of the ap- pearance of goosefoot is also problematic because C. album’s ability to germinate occurs during the entire growing season and this weed may yield seeds several times. Additionally, due to the fact that these seeds hold moisture well, even small numbers per 1 ha may disturb the long-term storage of grain. Determination of such weed thresholds is quite di�cult due to many factors that occur (e.g. the moment of weed appearance in relation to the developmental stage of the crop, weather and soil conditions and the actual prices of plant protection treatments) (Rola et al., 2013). However, research conducted by Valizadeh and Mirshekari (2011) suggests that the C. album threshold for rapeseed may be 4–8 pcs per m2 (coverage of rapeseed in the terms of LAI drops from 3 to 1.5). �e lack of selective herbicides, which at the same time could control the occurrence of goosefoot and be harm- less to plants (Kucharski, Sekutowski, 2007) means that only regular mechanical treatments help in reducing the occurrence of goosefoot in crop �elds. If neces- sary, chemical weed control is used along with areas adjacent to them, e.g. wasteland, ditches, paths and areas prepared for growing plants. It is also important to clean the equipment used for sowing seeds and harvesting plants, fertilise the soil only with well-spread manure and use good quality seed that is free from weed seed contamina- tion (Domagała-Świątkiewicz, 2007). Although the controversy around the phenomenon of allelopathy has not yet been unequivocally resolved, the negative e�ect of one plant on other neighbouring indi- viduals remains an indisputable fact (Gniazdowska, 2007). WesTon (2005) empha- sised that all plants contain allelochemical substances with di�erent structures and functions. �e germination and early growth analyses carried out have revealed the allelopathic e�ects of C. album extracts at various concentrations, which similarly interfere with the metabolism of grains and seeds and indicate the need for further research. Poonia et al. (2015) emphasized that attention should be paid to C. album not only as a bothersome weed but as a source of functional nutrients and useful me- dicinal properties. However, this species is a good host for many dangerous crop pests (Agrotis segetum Denis & Schi�ermüller, Aphis fabae Scop., Delia radicum L., Het- erodera schachtii Schmidt, Silphida latreille Latreille) and the host of some pathogens that cause viral diseases (e.g. beet mosaic, hepatitis jaundice (Abe, Tamada, 1986). C. K at ar zy na L ip ni ak , A ng el ik a K lis zc z 120 album was also reported to be the host of a  new plant disease caused by the fungus Stagonospora atriplicis (Westend.) Lind in New Zealand (McKenzie, Dingley, 1996). Conclusion (1) �e results presented in this paper con�rm the allelopathic properties of aqueous extracts of dry goosefoot (Chenopodium album). Based on the observations, an inhib- itory e�ect of allelopathic compounds released from C. album shoots on the GIs of triticale grains (×Triticosecale) and radish seeds (Raphanus sativus) was found. (2) Biometric analysis of triticale and radish seedlings demonstrated a signi�cant inhi- bition of growth in the presence of aqueous extracts from dry C. album shoots. (3) With an increasing concentration of C. album extracts, the FM values of 7-day trit- icale and radish seedlings were decreased. �e DM and the percentage of water con- tent values changed, depending on the type of seed and the concentration of goosefoot shoot extract. (4) �e negative e�ect of C. album was con�rmed by the electrolyte leakage detected from triticale and radish seedlings cells. With increasing allelopathic substances in the shoot extracts, an increase in water-ion balance disorders in the studied seedlings was observed. Con�ict of interest �e authors declare no con�ict of interest related to this article. References Abe, H., Tamada, T. (1986). Association of beet necrotic yellow vein virus with isolates of Polymyxa betp Keskin. 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Percentage of germinated triticale grains (×Triticosecale Wittm. ex A.Camus) – (A) and radish seeds (Raphanus sativus L.) – (B) watered with the aqueous extracts of Chenopodium album L. shoots in di�erent concentrations (0.5%, 1.0%, 1.5%, 2.0%) and control conditions Aqueous extracts [%] Days [24h] 1 2 3 4 5 6 7 A B A B A B A B A B A B A B Control 80 a ±1.58 80 a ±0.84 96 a ±0.89 84 a ±0.84 96 a ±0.71 96 a ±1.14 96 a ±0.55 100 a ±0.55 100 a ±0.00 100 a ±0.45 100 a ±0.00 100 a ±0.45 100 a ±0.00 100 a ±0.00 0.5 68 ab±0.71 60 ab ±1.14 80 ab ±0.89 64 ab ±1.30 96 a ±2.07 80 a ±2.28 96 a ±1.64 92 a ±1.92 100 a ±0.00 96 a ±1.30 100 a ±0.00 100 a ±0.45 100 a ±0.00 100 a ±0.45 1.0 40 b±1.14 8 c ±1.14 44 b ±0.84 12 b ±1.48 64 b ±2.97 16 b ±1.52 68 b ±2.07 20 b ±1.52 72 b ±1.52 28 b ±1.52 76 b ±1.22 44 b ±2.41 80 b ±1.30 44 b ±2.49 1.5 36 b±1.14 0 c ±0.00 40 b ±0.84 8 b ±0.71 56 b ±2.07 8 c ±1.14 60 b ±2.59 16 b ±0.84 64 b ±2.59 28 b ±1.14 68 b ±2.30 32 bc ±0.89 76 b ±2.17 40 b ±0.84 2.0 20 c±1.14 0 c ±0.00 28 c ±1.30 8 b ±1.30 52 b ±3.05 8 c ±1.14 56 bc ±2.68 16 b ±1.14 56 b ±2.97 20 bc ±1.48 60 bc ±3.85 24 c ±2.07 60 bc ±3.85 28 c ±2.07 mean values ±SD (n = 3) marked letter (a, b, c) in columns di�er signi�cantly according to Duncan’s test at p < 0.05 Tab. 2. �e germination parameters of triticale grains (×Triticosecale Wittm. ex A.Camus) – (A) and rad- ish seeds (Raphanus sativus L.) – (B) watered with the aqueous extracts of Chenopodium album L. shoots in di�erent concentrations (0.5%, 1.0%, 1.5%, 2.0%) and control conditions Aqueous extracts [%] CRG T50 GI SE A B A B A B A B Control 24.34 a 24.12 a 0.35 b 0.35 cd 58.74 a 59.42 a 80.00 a 80.00 a 0.5 23.70 a 22.98 ab 0.59 ab 0.67 a 53.74 a 55.42 ab 68.00 b 60.00 b 1.0 22.56 ab 19.72 b 0.67 a 0.59 ab 34.71 b 36.30 b 50.00 c 18.18 c 1.5 22.42 ab 18.44 bc 0.66 a 0.63 a 31.16 b 32.57 b 47.37 d 0.00 d 2.0 21.90 b 19.12 b 0.65 a 0.40 c 23.78 c 27.17 c 33.33 e 0.00 d the germination index (GI), speed emergence (SE), the coe�cient rate germination (CRG) index and the time needed to reach 50% of germinated seeds (T50); mean values (n = 3) marked letter (a, b, c) in columns di�er signi�cantly according to Duncan’s test at p < 0.05 Tab. 3. Seed vigour index (SVI) triticale (×Triticosecale Wittm. ex A.Camus) – (A) and radish (Raphanus sativus L.) – (B) seedlings a�er 7 days of germination on the aqueous extracts of Chenopodium album L. shoots in di�erent concentrations (0.5%, 1.0%, 1.5%, 2.0%) and control conditions SVI Concentration of aqueous extracts [%] Control 0.5 1.0 1.5 2.0 A B A B A B A B A B Underground organ 98.40 a 96.00 a 90.00 ab 64.80 b 53.25 b 10.91 c 39.74 c 3.50 d 5.67 d 12.50 e Aboveground organ 114.40 a 49.60 d 99.60 b 80.60 b 76.75 c 72.27 c 95.26 b 19.50 e 23.67 d 6.07 f Whole seedlings 212.80 a 145.60 a 189.60 b 145.40 a 130.00 c 83.18 b 135.00 c 23.00 c 29.33 d 18.57 d mean values (n = 3) in row marked with letters a, b, c di�er signi�cantly according to Duncan’s test at p ≤ 0.05 A llelopathic effect of goosefoot on germ ination and early stage grow th of triticale and radish 125 Tab. 4. Length and inhibition percentage (IP) of growth (expressed as a percentage of control) of triticale (×Triticosecale Wittm. ex A.Camus) and radish (Raphanus sativus L.) seedlings germinated on the aque- ous extracts of Chenopodium album L. shoots in di�erent concentrations (0.5%, 1.0%, 1.5%, 2.0%) and control conditions Organs Concentration of aqueous extracts [%] Control 0.5 1.0 1.5 2.0 [mm] [mm] IP% [mm] IP% [mm] IP% [mm] IP% Triticale Underground parts 98.4 a ±1.90 90.0 a ±2.37 6.31 42.6 b ±0.52 55.89 30.2 c ±0.91 68.87 3.4 d ±0.15 96.64 Aboveground parts 114.4 a ±1.21 99.6 a ±1.65 12.97 61.4 b ±2.09 46.79 72.4 c ±1.36 36.76 14.2 d ±0.80 87.35 Whole seedlings 212.8 a ±2.52 189.6 a ±3.83 104.20 104.0 b ±2.10 51.15 102.6 b ±2.14 51.72 17.6 c ±0.79 91.57 Radish Underground parts 49.6 a ±4.15 6.48 b ±0.92 –9.06 4.8 c ±0.87 93.83 1.4 d ±0.30 98.14 3.5 c ±0.29 94.39 Aboveground parts 96.0 a ±0.43 8.06 c ±2.03 –64.20 31.8 b ±0.30 35.85 7.8 c ±0.09 84.45 1.7 d ±0.17 96.51 Whole seedlings 14.56 a ±4.43 14.54 a ±2.88 –19.25 3.66 b ±0.93 72.55 0.92 c ±0.38 93.21 0.52 c ±0.19 96.07 minus (–) values of IP indicates growth stimulation, and plus (+) values of IP indicates growth inhibition; mean values (n = 3) in row marked with letters a, b, c di�er signi�cantly according to Duncan’s test at p ≤ 0.05 K at ar zy na L ip ni ak , A ng el ik a K lis zc z 126 Appendix 2 Fi g. 1 . R ad is h se ed lin gs (R ap ha nu s s at iv us L .) on th e 3r d ( A –E ), 5t h ( F– J) a nd 7 th (K –O ) d ay o f g er m in at io n w at er ed w ith a qu eo us ex tr ac ts o f g oo se fo ot (C he no po di um a lb um L .) sh oo ts a nd d is til le d w at er : A , F , K – c on tr ol (d is til le d w at er ); B, G , L – 0 .5 % e xt ra ct ; C , H , M – 1 % e xt ra ct , D , I , N – 1 .5 % e xt ra ct ; E , J , O – 2 % e xt ra ct (P ho to . K . L ip ni ak ) A llelopathic effect of goosefoot on germ ination and early stage grow th of triticale and radish 127 Fi g. 2 . T ri tic al e se ed lin gs (× Tr iti co se ca le W itt m . e x A .C am us ) o n th e 3r d ( A –E ), 5t h ( F– J) a nd 7 th (K –O ) d ay s o f g er m in at io n w at er ed w ith a qu eo us e xt ra ct s o f g oo se fo ot (C he no po di um a lb um L .) sh oo ts a nd d is til le d w at er : A , F , K – c on tr ol (d is til le d w at er ); B, G , L – 0. 5% e xt ra ct ; C , H , M – 1 % e xt ra ct ; D , I , N – 1 .5 % e xt ra ct ; E , J , O – 2 % e xt ra ct (P ho to . K . L ip ni ak ) K at ar zy na L ip ni ak , A ng el ik a K lis zc z 128 Abstract �e aim of this study was to investigate the e�ect of aqueous extracts from Chenopodium album L. on ger- mination and early stages of triticale grains (×Triticosecale Wittm. ex A.Camus) and radish seeds (Raphanus sativus L.). Germination indexes, fresh and dry mass, water content and electrolyte leakage were measured. Studies revealed the di�erent germination capacity of triticale grains and radish seeds, where increased concentrations of allelopathins in aqueous C. album extracts signi�cantly inhibited seedling growth for both species. �e extracts had an inhibitory e�ect on the growth of seedling fresh mass. An increase in dry mass of radish seedlings was demonstrated for each of the extracts and, for triticale seedlings, only at concentrations of 0.5% and 1.5%. Water content in triticale and radish seedlings varied depending on the concentration of allelopathins in the extract. With increasing concentrations of C. album extract, regardless of seedling type, a statistically signi�cant increase in electrolyte leakage was observed. Key words: fresh and dry mass, seed germination indexes, electrolyte leakage, relative water content Received: [2020.03.16] Accepted: [2020.05.25] Allelopatyczny wpływ komosy białej na kiełkowanie i wczesne stadia wzrostu pszenżyta i rzodkiewki Streszczenie Celem pracy było zbadanie wpływu wyciągów wodnych z Chenopodium album L. na kiełkowanie oraz wczesne stadia wzrostu ziarniaków pszenżyta (×Triticosecale Wittm. ex A.Camus) i nasion rzodkiew- ki (Raphanus sativus L.). Wyznaczono wskaźniki kiełkowania, świeżą i suchą masę, zawartość wody oraz wyciek elektrolitów. Badania wykazały różną zdolność kiełkowania ziaren pszenżyta i  nasion rzodkiewki – wraz ze wzrostem stężeń substancji allelopatycznych zawartych w wodnych ekstraktach z  C. album obserwowano znaczne zahamowanie wzrostu siewek obydwu gatunków. Wodne wyciągi hamowały przyrost świeżej masy siewek. Natomiast przyrost suchej masy siewek rzodkiewki wyka- zano przy każdym z ekstraktów, a dla pszenżyta jedynie w stężeniach 0,5% i 1,5%. Zawartość wody w siewkach pszenżyta i rzodkiewki była zróżnicowana, w zależności od stężenia allelopatin w ekstrak- cie. Wraz ze wzrostem stężenia ekstraktów C. album, niezależnie od rodzaju siewek, zaobserwowano statystycznie istotny wzrost wycieku elektrolitów. Słowa kluczowe: świeża i sucha masa, wskaźniki kiełkowania nasion, wyciek elektrolitu, względna zawartość wody Information on the authors Katarzyna Lipniak She is interested in weeds in�uence on germination and growth of seeds. Angelika Kliszcz https://orcid.org/0000-0002-1270-4414 She is focusing on enhancing the understanding of the in�uence of di�erent factors on soil structure and fertility. Particularly, she is investigating the interaction of plants with the physical, chemical, and biological properties of the soil. She is also interested in earthworm ecology and mesofauna function in agroecosystems.