203 Annales Universitatis Paedagogicae Cracoviensis Studia Naturae, 6: 203–225, 2021, ISSN 2543-8832 DOI: 10.24917/25438832.6.12 Angelika Kliszcz Department of Agroecology and Plant Production, Faculty of Agriculture and Economics, University of Agriculture in Kraków, Al. Mickiewicza 21, 31-120 Kraków, Poland; angelika.kliszcz@urk.edu.pl Phenological growth stages and BBCH-identification keys of Jerusalem artichoke (Helianthus tuberosus L.) Introduction �e growth stages of development of particular plant species were constructed and pro- posed by many authors around the world since decades. Chen (2013) indicates that the tradition of observation and recording the phenological events of many cultivated and ornamental plants in ancient times were occurred. �e interesting plant species that is being rediscovered today is Jerusalem artichoke, topinambour (Helianthus tuberosus L.). Both, the growing attention of scientists in the context of its interesting physiological, biochemical and genetic predispositions invasive plant (Balogh, 2008; Tokarska-Guzik et al., 2012), resistant to salt stress (Zhang et al., 2016), quite di�cult to correctly iden- tify H. tuberosus and H. strumosus L. and their natural hybrids – they both produce tubers from all the other species of the Helianthus genus (Kays, Nottingham, 2008), as well as the growing interest in this plant as a raw material in the food industry (high inulin content, an easily hydrolysable fructan (Barhatova et al., 2015), component of sophisticated alcoholic beverages (Rossini et al., 2016), health-promoting properties of tubers (Cieślik, Filipiak-Florkiewicz, 2000; Kulczyński, Gramza-Michałowska, 2016; Radovanovic et al., 2014), make this plant nowadays more and more noticeable. Ad- ditionally, the energy sector approves this plant as an energetic crop (bioethanol and biofuel production, pellets production due to a large amount of biomass produced per unit area with a  satisfactory caloric value (Kowalczyk-Juśko et al., 2012; Johansson et al., 2015; Sawicka et al., 2019; Bedzo et al., 2020). �ere are also other applications (such as blasting in forest frames as an alternative food for wild animals (Dreszczyk, Brzezowska, 2008), especially wild boars, a  substrate medium for the production of mushrooms and shunts (Đorđević et al., 2010), or for industry processes with the use of microorganisms laboratory cultures (Meng et al., 2021) make this plant more and more famous. A ng el ik a K lis zc z 204 However, each multi-purpose plant has its limitations, among which it should be noted: invasive nature (high inulin content, thanks to which the tubers winter in the soil down to –30°C, rapid growth and early growth of the aboveground parts shading the surface, vegetative reproduction mostly climatic zones, remains in the position despite herbicidal fallow (Balogh, 2008; Tokarska-Guzik et al., 2012; Pacanoski, Mehmeti, 2020). �e increasing number of papers according Jerusalem artichoke in Web of Science (from 1910 to 2000: 615 articles, 2001–2010: 221 articles, 2011–2015: 292 articles, 2016–2020: 480 articles) and agricultural events focusing on Jerusalem artichoke (Stapor, 2020) is gathering more and more people interested in this plant around the world. More information about origin, history of discover and various naming of this plant can be found in �e Biology and chemistry of Jerusalem artichoke (Kays, Nottingham, 2008). Detailed information about probiotic and pharmaceutical properties of this plant was published by Mystkowska et al. (2015), its multipurpose use was provided by Sawicka et al. (2012), and energy properties of the plant announced Gao et al. (2016). Recently, the very valuable review, with a whole view on this plant appeared (Rossini et al., 2019). �e aim of the study is to focus on the growth and development of Jerusalem artichoke (H. tuberosus) plants grown from the tubers in temperate climate zone and propose a BBCH (Biologische Bundesanstalt, Bundessortenamt and Chemicsche In- dustrie) identi�cation key, which is expected for uni�cation academic and practical discussion about this plant. Due to the fact that in our climate the seeds are not fully developed and their role in propagation of the plant is negligible, the proposed BBCH key describes all development phases except the ripening of seeds phase (is omitted). �e developmental biology of Jerusalem artichoke – state of the art �e general strategy of Jerusalem artichoke (JA) is to invests actual carbon and nutri- ents early in its development into stem (Incoll, Neales, 1970), branch, and leaf growth, facilitating the exploitation of aboveground resources. Later in the developmental cycle carbon and nutrients are allocated to rhizomes and tubers (McLaurin et al., 1999), enabling the species to spread alongside, i.e. colonising new areas. �e success of its allocation patterns is in simultaneous synchronization of the below- and aboveground biomass growth and development (Fig. 1). �e plants produce considerable amounts of aboveground biomass, which acts for carbohydrates reservoir although it has C3 photosynthetic mode (Podlaski et al., 2017). Helianthus tuberosus plants have a strong photoperiod response (short-day plant) (Terzić et al., 2012). One of the �rst research exploring photoperiod phenomenon in plants was based on Jerusalem artichoke species (Garner, Allard, 1923). Shortly, photo- periodic plants identify day length in the leaves and then transfer the signal (�origen) to the shoot apex for the onset of the formation of in�orescence. �e day length also a�ects the start of formation tubers (i.e. tuberisation). Typically initiation of tuberisation 205 begins from 5 to 13 weeks a�er emergence (Swanton, Cavers, 1989; Hay, O�er, 1992; McLaurin et al., 1999). Generally, the development of aboveground biomass goes through successive phases, from the sprouting, full side branching (BBCH 39n), to full drying o� the plant (BBCH 98). �e belowground development starts with roots development (BBCH 04), then rhizome development (BBCH 40-49), and tuber development (BBCH 70-79). �e BBCH system of monitoring physiological scales in plants Any lively discussion between people from di�erent regions needs a common language. Except for the known need to communicate in the same language, it is nonetheless necessary to think about the same phenomena in the same way, especially when the discussion concerns a dynamic process in biological systems, additionally modi�ed by environmental processes in�uenced in various intensity in every point on the world map. Scientists’ passion, farmer’s needs, and entrepreneurs’ interests underlie the universal BBCH scale (Biologische Bundesanstalt, Bundessortenamt and Chemische Industrie) currently in force (Meier, 2018). �e Monograph did not appear at once (Meier et al., 2009). Many interesting works on this topic (Troitzky, 1925; Fleckinger, 1948; Feekes, 1941; Large, 1954) were published through the 20th century, to meet the demands of 1) Fig. 1. General view on developmental stages of Jerusalem artichoke (Helianthus tuberosus L.) P henological grow th stages and B B C H -identification keys of Jerusalem artichoke (H elianthus tuberosus L.) A ng el ik a K lis zc z 206 uni�cation and clari�cation of de�nitions of concepts in botanical scienti�c discussion, 2) simpli�cation the decision making in process of plant protection by farmers and avoid the misunderstanding between farmers and agrochemical companies or agricultural insurance agents, and 3) development of agrometeorology. However, a real acceleration of universal concepts describing phenological stages during plant development occurred a�er a publication scale by Zadoks et al. (1974). �e authors presented an adjusted and re�ned numeric decimal scale for such plants like cereals and rice, which gave the direct base for currently wide-spread used BBCH scale. �e BBCH coding system is an improvement of the Zadoks et al. (1974) coding system, it includes also the dicotyledoneous plants and more monotyledoneous plants species. �e �rst publication of the BBCH codes of some crops (Bleiholder et al., 1989) (appears in working group consisted of sta� members from four chemical companies) was the �rst step to join the forces in 1991 with German scientists (from �e Federal Biological Research Centre for Agriculture and Forestry, BBA), who published booklets describing phenological stages of particular crops (Meier, 1985). �e �rst outcome of this cooperation was the principles of the enhanced general BBCH scale (Hack et al., 1992), which was the base for members of this group for publishing (with experts in each crop) the “extended BBCH scales for speci�c crops” in various branch journals. �e �rst BBCH Monograph edited by Meier (1997) was published in four languages and describes the phenological development stages of 27 crops and wild plants. Adam- czewski and Matysia (2005) published the BBCH scale in Polish, a�er earlier published work (e.g. Gąsowski, Ostrowska, 1993). Milestone for the international acceptance of the BBCH codes used in plant protection management process was the decision for establishing the BBCH scale mandatory for all o�cial plant protection trials made in 2004 and 2006 by EPPO (European and Mediterranean Plant Protection Organization) (Meier et al., 2009). Nowadays the BBCH Monograph (Meier, 2018) includes 48 identi�cation keys for crops and additional key for weeds (Dicotyledons, Graminae, Monocotyledons, Perennial plants). Recently, the developmental biology of many other crops has been described in a key on the BBCH scale, e.g. to harmonize production processes (Rajan et al., 2011; Zhao et al., 2019; Singh et al., 2021). Although, despite of multi-purpose use and increased awareness of H. tuberosus species, there is a lack of o�cial BBCH identi�cation key describing phenological growth stages of H. tuberosus. In this article, developmental biology of Jerusalem artichoke in temperate climate was presented. �e phenological stages of this plant were proposed in the obligatory BBCH identi�cation key. 207 Material and methods Experimental site and plant material �e plants were grown in experimental �eld located in Experimental Station in Krakow – Mydlniki (50°05′08.5″N; 19°51′08.3″E; University of Agriculture in Kraków, Poland) in 2020. During the 7 months (mid-April to mid-November 2020) growth and devel- opment of Helianthus tuberosus plants (cv. Rubik) were monitored. �e seed tubers were intentionally le� in the �eld previous autumn (November 2019) as an irregular population close to the natural one. In spring, plants were chosen and monitored dur- ing the vegetation season (completely random assignment was applied). �e soil was unfertilized and characterised a well moisture content during whole vegetation period (due to impact of an neighbourhood of underground watercourse). �e BBCH scale �e BBCH principal growth stages were the basis for these considerations (Tab. 1). Tab. 1. Principal growth stages of Helianthus tuberosus L. according to BBCH scale (a�er Meier et al., 2009) Stage (number 0–9) Description 0 Germination / sprouting / bud development 1 Leaf development (main shoot) 2 Formation of side shoots/tillering 3 Stem elongation or rosette growth/shoot development (main shoot) 4 Development of harvestable vegetative plant parts or vegetatively propagated organs / booting (main shoot) 5 In�orescence emergence (main shoot) / heading 6 Flowering (main shoot) 7 Development of fruit 8 Ripening or maturity of fruit and seed 9 Senescence, beginning of dormancy Meteorological data �e observations were carried out with the background of weather conditions in tem- perate climate (Poland). �e meteorological data was obtained from ClimateData.org (2020). �e graphical relation of mean temperatures [°C] and total precipitation values [mm] for each month were reported as a Gaussen-Walter climatogram with Łukasiewicz modi�cation (Walter, 1976; Łukasiewicz, 2006) (Fig. 2). �e rule for the construction of such a graph is that the values of mean temper- ature and total precipitation are plotted with maintaining a  ratio of 1°C to 4 mm of precipitation. �is balance determines the di�erence between precipitation and evap- otranspiration. It helps to read o� the amount of evapotranspiration and thus estimate the excess or shortage of precipitation for plants on a local scale (Treder et al., 2018). P henological grow th stages and B B C H -identification keys of Jerusalem artichoke (H elianthus tuberosus L.) A ng el ik a K lis zc z 208 Results �e stages of phenological phases of Helianthus tuberosus was constructed for temperate climate zone (Tab. 2 – Appendix 1). Principal growth stage 0: sprouting Helianthus tuberosus tubers have endodormancy, which means that an internal mech- anism prevents sprouting even though the environmental conditions may be suitable. It is linked with the conditions (winter months) of their region of origin (temperate zone, continental climate). �e main shoot develops from apical bud located on the belowground seed tuber (Fig. 3A–B). �ere are possible other lateral sprouts growing from the same tuber at the same time (from axillary buds), but the plant seems to develop lateral sprouts a little bit later so as not to inhibit the growth of the main shoot. Because some tubers are located fairly deep in the ground, the length of the �rst sprout may achieve even 30 cm (BBCH 08, Fig. 3B), due to high vitality and aÈuence of substances in tubers. Sprouts a�er reaching a suitable cumulative temperatures grow in the soil quite fast regardless of weather conditions. Fig. 2. Gaussen-Walter climatogram for the studied area (in months January–November 2020) 209 Principal growth stage 1: leaf development (main shoot) �e main shoot (stem) grows quite fast and tends to shade the stand. �e development of aboveground leafy biomass realizes quickly. When the plant meets favourable weather, it generates next leaves pair in each new week. Leaves are numerous, with the opposite arrangement in the lower third, alternate above. In the studies about the dynamics of this stage, it will be worth to mark with an asterix the last pair of the leaves with opposite arrangement (e.g. BBCH16*). Generally, the number of leaves pairs depend on biotope richness, and to a lesser extent on the weather conditions. Once the side branches appear from the apical buds in any leaves pair, the development of the plant shall be determined in accordance with appropriate next phase (BBCH 3_). Principal growth stage 2: formation of other sprouts �is stage occurs not always here. Generally, the plant skips with its development from leaf development phase (BBCH 1_) to the next phase (BBCH 3_). �e strategy of for- Fig. 3. �e examples of some phenological stages of proposed BBCH coding system for Jerusalem artichoke (Helianthus tuberosus L.) (Photo. A. Kliszcz) P henological grow th stages and B B C H -identification keys of Jerusalem artichoke (H elianthus tuberosus L.) A ng el ik a K lis zc z 210 mation of other stems (lateral sprouts) from the underground parts of the plant (tuber, rhizomes, underground part of the main stem) seems to be linked with the need for more photosynthetic area within the plant, what constitutes their carbohydrate supply for new tubers or it is a result of mechanically injured belowground stem. �e number of shoots that emerge are linked with e.g. shape of tuber and it is variety depended. �e tuber more branched produces more shoots (tubers branched vs. tuber with apical dom- inance) (Kays, Nottingham, 2008). �e formation of other sprouts from the tuber also depends on the depth at which the tuber is located, tuber size and number of its axillary buds. In general, since the delineation of the sources of additional stems is unclear, all shoots that came above the ground surface a�er the main shoot drop into one category. It is worth to notice that the emergence of other sprouts (stems) may appear during the whole vegetative phase. �e �nal number of stems is constituted at the end of the growing season, when the plant enters the senescence phase. Principal growth stage 3: stem elongation and development of lateral branches on main shoot (side branching) In this stage the development of the stem and upper leaves continues. Side branching appears simultaneously on the main shoot from the bottom of the plant. �e presence and degree of branching depend on the variety, plant population density, and other factors (like branch location on the plant, photosynthetic potential, environmental factors). �e lateral branches are formed in the axils of the leaves, starting at the base of the plant. At each node, there is commonly two opposite-located lateral branches. Very rare is the triple, when three leaves emerge from one node. Rapid growth of branches on the plant diminishes in the middle of growth cycle and again increases when axillary buds start to developing into �owering branches. Vertical development of the plant is terminated by �ower bud formation at the apex of the stems. Not all BBCH codes may occur during the development of the plant in this stage. As soon as the next axillary buds begin to develop in the next (upper) pair of leaves on the plant, the BBCH code is counted there (e.g. BBCH 32-1 move to BBCH 33-0, even if the leaf development on the lower branch continues). �e size of axillary buds should be at least 2 cm to be considered as the next level of this development phase (Fig. 3D). Typically, every next node with leaves pair generates the twin opposite branches, but it depends on the plant’s current needs (e.g. shading, processes of translocating carbohy- drates). �e plant may omit some pairs of leaves without developing any branches there. If this is the case for the third pair of leaves, the BBCH 33-0 remains until a fourth pair of leaves has developed and the axillary buds in the axils of this leaves appear. Another case may arise when fourth pair of leaves has already hosted emerging twin branches (e.g. already with two leaves each), then the BBCH 33-_ is valid until in the axis of the ��h pair of leaves appear new axillary buds. 211 Principal growth stage 4: rhizomes booting �e plant starts to form rhizomes in the early stages of biomass acquisition and accumu- lation (from 1.5 to 8 weeks a�er emergence; Fig. 3E). �e underground portion of the stem (4 to 5 cm below the soil surface) is the basis for the emergence of rhizomes. �ey grow in a slight downward angle (Fig. 3F) with internodes length varying substantially among clones (Kays, Nottingham, 2008). Principal growth stage 5: in�orescence emergence �e shi� between vegetative and generative phase in Jerusalem artichoke (short-day clones) is strongly dependent on photoperiod. �e in�orescence appears in the speci�c location on the plant of JA (i.e., the top of the stem and later – on branch apices) and their formation involves temporal order. Because of that the BBCH coding system for this phase was focused on �rst ap- peared examples of top in�orescence, and then on branched-located ones (e.g. BBCH 51 -> BBCH 52, 53). Long-cycle varieties (ca. 9 months vegetation) o�en produce buds but no �owers (Denoroy, 1996), and their aerial parts are moving then straight on to senescence phase (BBCH 9_). Principal growth stage 6: �owering �e �owering is starting from the top of the main shoot (BBCH 61-65) and will proceed to the bottom (along with side shoots embedded on the main shoot, i.e. BBCH 66-69). Flower stalks have frequently between 10 and 15 cm for Rubik cultivar, depending on closeness to the plant axis. �is paper focus on domesticated clone (cv. Rubik), and it was observed that in temperate climate (Fig. 2) it takes ca. 21 days for the plants to proceed �owering (from the tight bud stage (BBCH 51) to senescence stage (BBCH 90)). �e detailed visualisation of chronological sequence of �owering phase of single �ower was presented in the book concerning the biological and chemical issues of JA plants (Kays, Nottingham, 2008). �e BBCH scale required the �owering sequence of all �owers on the plant due to the fact that JA plants produce many �owers (not a  single one as the sun�ower produces). And for that reason presented BBCH scale includes the whole �owering biology of this plant with the background of single �ower development steps. Principal growth stage 7: tuber bulking While tuber initiation appears to be in part controlled by carbohydrate supply, tuber bulking is strongly modulated by photoperiod, even in clones that are day neutral for �owering (Kays, Nottingham, 2008). Cumulative temperature is linearly correlated with tuber number, and cumulative degree days (≥ 520 degree days) can be used for predicting the onset of tuberisation (Spitters et al., 1988). In the cross section tuber P henological grow th stages and B B C H -identification keys of Jerusalem artichoke (H elianthus tuberosus L.) A ng el ik a K lis zc z 212 could be divided (from exterior to interior) into: epidermis, cortex, outer medulla, inner medulla, and pith (Mazza, 1985). Shortly a�er the beginning of �owering, remo- bilization of nutrients from canopy into the developing tubers begins. Photoperiod and carbohydrate supply are a critical factors in the tuberisation response. Simultaneously with the proceeding bulking stages of the tuber, the dormancy onset occurs. �e onset of dormancy is progressing gradually. Firstly, into dormancy enter rhizomes and small, young tubers, with more and more areas of the tuber establishing dormant. �e larger, more mature tubers enter into dormancy at the end within the whole plant. Although, the initiation of dormancy in large tuber may occur even the tuber is not fully �lled (Kays, Nottingham, 2008). Principal growth stage 8: ripening seed Flowers are o�en sterile (domesticated clones). Swanton et al. (1992) stated that most of the JA plants have no more than 5 seeds per �ower head. According to Westley (1993) only a  44 % of mature seeds are capable of germination, whereas only 33% are able for reproduction – during the �rst season (wild clones). In this phase, the increased translocation of the assimilates to the tubers takes place, and this coincides with and a�ect the seed ripening as well. Because the propagation of the plant by the seed is of marginal importance, and it occurs when the plant is drying o� (transfer of the sym- bionts to the tubers, i.e. tuber bulking, Fig. 3G–H), no special phase was assessed for process regarding seeds ripening. Principal growth stage 9: senescence �e drying o� of the whole plant occurs when the belowground part of the plant enters dormancy. �e process of senescence accompanies the plant virtually throughout the entire growing season. �e �rst leaves on the main stem senesced �rstly, and it happens well before it starts blooming. Some authors argue this fact that the plant shades the lower leaves as it grows (Zubr, 1988). �e meteorological data (Fig. 2) shows no shortages of water in the studied period, i.e. the precipitation line is above the line representing the temperature, Such ratio (1°C to 4 mm precipitation) is proposed by Łukasiewicz (2006) for climate conditions in Poland (temperate). �erefore, the plants had favourable conditions for development during whole vegetation season. Discussion �e Jerusalem artichoke (Helianthus tuberosus L.) is an aged tuber crop with a lately aroused attraction following its multipurpose usage (Cao et al., 2008; van Wyk, Wink, 2008; Ma et al., 2011; Maj et al., 2013; Mystkowska, Zarzecka, 2013). �e growth of 213 plants and certain developmental phenomena are governed by a few main factors: the total amount of heat a plant received during a certain period, the portion of various wavelengths from sunlight, number of days with sunlight and the duration of the day, richness of soil and their su�cient moisture (Biggs et al., 2007; Kocsis et al., 2007; Puangbut et al., 2012; Ruttanaprasert et al., 2014). However, the developmental biology of wild, domesticated, and intermediate clones di�er signi�cantly, but general overview of phenological stages of Jerusalem artichoke seems to be constant (Fig. 1). �e plant have to go through all stages to produce tubers (Fig. 3A), a main propagules for next growing season. However, the development of this species is realised in two parallel directions: vegetative and generative (Pawłowski, Jasiewicz, 1971; Vaughan, Geissler, 2001). �is complicates the naming of its developmental stage. Dual naming can be a  solution. When the side branching phase continues, the rhizomes booting occurs, and then the proper description could be e.g. BBCH 38-2/46. Similarly, as the tubers �ll, the aboveground biomass withers (it could be written as BBCH 78/97). Of course, the codes can be used individually to indicate only the state of the aerial vegetative biomass (respectively, BBCH 38-2, or BBCH 97). But for detailed ecological studies of this plant it is bene�cial to use dual nomenclature. �is paper focuses on domesticated clone (cv. Rubik), which belongs to the group of short-day clones. �e �rst attempt to name and standardise Jerusalem artichoke (JA) developmental biology stages was made by Paungbut et al. (2015). However, they designed their own system without using the BBCH codes. �ey arrange all development of JA into three main groups of stages: Vegetative stages (V), Reproductive stages (R), and Tuberisation stages (T). �e authors cultivated the plants in �ailand (tropical area) in 2011–2012, and observed developmental biology of JA during opposite seasons, both, the early-rainy season and the drier post-rainy season. �ere are 15 phases in their concept (adding to this number of pairs of leaves on the main stem, depending on the nodes produced; shortly, if the main stem develops 12 leaves pairs, there will be 25 phases in total). �e BBCH codes allow for a more accurate description of each phase through careful observation of plant morphology. �erefore, in presented concept according to BBCH codes there are at least 100 various phases, which precisely de�ne every moment of development (e.g. BBCH 3_ is describing side branching). More detailed descriptions of the JA plants are already included in the phenotype studies of the various clones of this species (Diederichsen, 2010), which is no necessary to include it in BBCH system formation. It is worth to note that with modi�ed biotope features (unfavourable conditions), the length of developmental stages could be shorten (e.g. the time of bulking tubers) or abandoned (the �owering phase does not always occur). Additionally, the genetic features play a pivotal role as well (Kays, Kultur, 2005; Skiba, Sawicka, 2016) and also wild vs. domesticated clones develop with di�erent dynamics (Feher et al., 1999; Breton P henological grow th stages and B B C H -identification keys of Jerusalem artichoke (H elianthus tuberosus L.) A ng el ik a K lis zc z 214 et al., 2017). For example, Serieys et al. (2010) examined 142 clones of JA deposited in INRA library and stated that 80% of them perform the �owering phase between September and October, and 10% of them did not enter this phase at all. On the other hand, the biomass enhancement and its dynamics could be induced with richness of site or fertilisation level (Bogucka et al., 2021). Interesting research on the dynamics of nutrient uptake was carried out by Izsaki and Németh (2013). �e authors examined two varieties of JA and stated that in both cases the maximum nutrient uptake was recorded on the 155th day of vegetation, which corresponds with beginning of 7th phase (BBCH 70-79), i.e. tubers bulking. Manipulation of duration of development processes could be intentionally forced in agricultural practice, which is targeted on high-quality, plentiful tubers yield. In literature the manipulation of planting date (Puangbut et al., 2012), mowing date of plant top (Acar et al., 2011), dates of pruning radius (Gao et al., 2018), harvest time and storage (Saengthongpinit, Sajjaanantakul, 2005) or another agrotechnical treatments are known (Puttha et al., 2013; Dias et al., 2016; Gao et al., 2019). �erefore, the proper managing of the plantation of topinambur plants seems to be an important factor for high yield of tubers in many agroecosystems. Conclusion �e scale for coding the phenological growth stages of Helianthus tuberosus L. species is needed. �is plant becomes more and more popular because of its multi-purpose use and ease of cultivation in all types of soil all over the world. �e latest edition of BBCH Monograph (Meier, 2018) does not cover this need. Jerusalem artichoke has many desirable growing traits such as cold and drought tolerance, wind and sand resistance, saline tolerance, strong fecundity and high pest and disease resistant. Generally, the development of aboveground biomass goes through successive phas- es, from the sprouting, full side branching (BBCH 39n), to full drying o� of the plant (BBCH 98). �e belowground development starts with roots development (BBCH 04), then rhizome development (BBCH 40-49), and tuber development (BBCH 70-79). It is worth to notice that the development of the plant from a point (BBCH 49) realises in two parallel directions: vegetative and generative, i.e. when at least one rhizome starts to thicken at its end, it means that the plant begun a generative phase. �e concern about their developmental biology is also essential for managing the termination strategies for this genus, as the JA is an invasive plant in most ecosystems. �erefore growing attraction of this plant force their key in BBCH coding system to harmonize discussion about this plant in the future. 215 Acknowledgements �e research was �nanced by the Ministry of Science and Higher Education of Poland. Con�ict of interest �e author declares no con�ict of interest. References Acar, R., Ada, R., Özköse, A. (2011). e�ects of di�erent mowing dates of plant top on tuber yield of Jerusa- lem artichoke (Helianthus tuberosus L.). African Journal of Biotechnology, 10(45), 9036–9040. https:// doi.org/10.4314/ajb.v10i45. Adamczewski, K., Matysia, K. (2005). Klucz do określania faz rozwojowych roślin jedno- i dwuliściennych w skali BBCH (�e key to determining the development phases of monocotyledonous and dicotyledonous plants on the BBCH scale). 1st ed. Poznań: Instytut Ochrony Roślin. [In Polish] Amarowicz, R., Cwalina-Ambroziak, B., Janiak, M.A., Bogucka, B. (2020). E�ect of n fertilization on the content of phenolic compounds in Jerusalem artichoke (Helianthus tuberosus L.) tubers and their antioxidant capacity. Agronomy, 10(8), 1215. https://doi.org/10.3390 /agronomy10081215. Balogh, L. (2008). Sun�ower species (Helianthus spp.)’. In the most important invasive plants in Hungary. Hungary: Institute of Ecology and Botany, Hungarian Academy of Sciences, p. 227–255. Barhatova, T., Nazarenko, M., Koguhova, M., Hripko, I. (2015). Obtaining and identi�cation of inulin from Jerusalem artichoke (Helianthus tuberosus L.) tubers. Foods and Raw Materials, 3(2), 13–22. https://doi.org/10.12737/13115 Bedzo, O.K.K., Mandegari, M., Görgens, J. (2020). Techno-economic analysis of inulooligosaccharides, protein, and biofuel co-production from Jerusalem artichoke tubers: a biore�nery approach. Biofuels, Bioproducts and Biore�ning, 14(4), 776–793. https://doi.org/10.1002/bbb.2105 Biggs, M., McVicar, J., Flowerdew, B. (2007). Wielka księga warzyw, ziół i owoców (�e great book of vege- tables, herbs and fruits). Warszawa: Dom Wydawniczy Bellona, p. 110–111. [In Polish] Bleiholder, H., van den Boom, J., Langelüddecke, P., Stauss, R. (1989). Einkeitliche codierung der phänolo- gischen stadien bei kultur-und schadp�anzen (Uniform coding of the phenological stages in cultivated and harmful plants). Einkeitliche Codierung Der Phänologischen Stadien Bei Kultur-Und Schadp�anzen, 41(11), 381–384. [In German] Bogucka, B., Pszczółkowska, A., Okorski, A., Jankowski, K.(2021). �e e�ects of potassium fertilization and irrigation on the yield and health status of Jerusalem artichoke (Helianthus tuberosus L.). Agronomy, 11(2), 234. https://doi.org/10.3390/agronomy11020234 Cao, H., Liu, Q., Li, S., Zhao, Z., Du, Y. (2008). Helianthus tuberosus – A good kind of biomass source for dimethylfruran production. Journal of Biotechnology, 136(1), S271–S272. Chen, X. (2013). East Asia. In: M.D. Schwartz (ed.), Phenology: an integrative environmental science, p. 9–22. Dordrecht: Springer Netherlands. https://doi.org/10.1007/978-94-007-6925-0 Cieślik, E., Filipiak-Florkiewicz, A. (2000). Topinambur (Helianthus tuberosus L.) – możliwości wykorzy- stywania do produkcji żywności funkcjonalnej (Topinambur (Helianthus tuberosus L.) – the possibility of using it in the production of functional food). Żywność, 1(22), 9. [In Polish] ClimatData.org (2020). Kraków-Klimat (Polska). Available at: https://pl.climate-data.org/europa/polska/ lesser-poland-voivodeship/krakow-715022/#climate-table, Accessed on 26 May 2021. Denoroy, P. (1996). �e crop physiology of Helianthus tuberosus L.: a model orientated view. Biomass and Bioenergy, 11(1), 11–32. https://doi.org/10.1016/0961-9534(96)00006-2 Dias, N.S., Ferreira, J.F.S., Liu, X., Suarez, D.L. (2016). Jerusalem artichoke (Helianthus tuberosus L.) maintains high inulin, tuber yield, and antioxidant capacity under moderately-saline irrigation waters. Industrial Crops and Products, 94, 1009–1024. https://doi.org/10.1016/j.indcrop.2016.09.029. P henological grow th stages and B B C H -identification keys of Jerusalem artichoke (H elianthus tuberosus L.) A ng el ik a K lis zc z 216 Diederichsen, A. (2010). Phenotypic diversity of Jerusalem artichoke (Helianthus tuberosus L.) ger- mplasm preserved by the Canadian Genebank. Helia, 33(53), 1–15. https://doi.org/10.2298/ HEL1053001D. Đorđević, N., Popović, Z., Grubić, G., Vučković, S., Simić, A. (2010). Production of fodder in the hunting grounds for game feeding and decrease of damages in agriculture and forestry. Forage Crops Basis of the Sustainable Animal Husbandry Development, Biotechnology in Animal husbandry, UDC 636(26), 539–547. Dreszczyk, E., Brzezowska, J. (2008). Ocena przydatności roslin do uprawy alternatywnej z wykorzystaniem typowych systemów technicznych (Assessment of the suitability of plants for alternative cultivation with the use of typical technical systems). Materiały XIII Międzynarodowej Konferencji Naukowej „Inżynieria Rolnicza a Środowisko”, s. 102–103. Międzyzdroje. [In Polish] Feekes, W. (1941). De tarwe en haar milieu (�e wheat and its environment). Verslag Technische Tarwe Commissie 12, 523–888. [In German] Feher, A., Koncekova, L., Lisyova, J. (1999). Invasive behaviour in wild populations of Jerusalem Artichoke (Helianthus tuberosus). In: Slovenska Polnohospodarska Universita. Nitra, p. 79–82. https://agris.fao. org/agris-search/search.do?recordID=SK2000000315 Fleckinger, J. (1948). Les Stades Vegétatifs Des Arbres Fruitiers, En Rapport Avec Le Traitements (Vegetative stages of fruit trees, Related to treatments). Pomologie Française, Supplément, 81–93. Gao, K., Tie-Xia, Z., Lin, W., Yang, G. (2018). E�ects of root pruning radius and time on yield of tuberous roots and resource allocation in a crop of Helianthus tuberosus L. Scienti�c Reports, 8(1), 4392. https:// doi.org/10.1038/s41598-018-22586-1 Gao, K., Tie-Xia, Z., Qi-Bing, W. (2016). Nitrogen fertilization, irrigation, and harvest times a�ect biomass and energy value of Helianthus tuberosus L. Journal of Plant Nutrition, 39(13), 1906–1914. https://doi. org/10.1080/01904167.2016.1189938 Gao, K., Zhixin, Z., Tiexia, Z., Xun, T., Yang, G., Lixing, Z., , Tianqi, L. (2019). �e in�uence of leaf removal on tuber yield and fuel characteristics of Helianthus tuberosus L. in a semi-arid area. Industrial Crops and Products, 131, 8–13. https://doi.org/10.1016/j.indcrop.2019.01.024 Garner, W.W., Allard, H.A. (1923). Futher studies in photoperiodism, the response of the plant to relative length of day and night. Journal of Agricultural Research, 23(2), 871–920. Gąsowski, A., Ostrowska, D. (1993). Klucz do oznaczania stadiów rozwojowych niektórych gatunków roślin rolniczych (�e key to determining the development stages of some agricultural plant species). Warszawa: Wydawnictwo SGGW. [In Polish] Hack Von H., Bleiholder, H., Buhr, L., Meier, U., Schnock-Fricke, U., Weber, E., Witzenberger, A. (1992). Einheitliche Codierung der phanologischen Entwicklungsstadien mono- und dikotyler P� anzen – Erweiterte BBCH-Skala, Allgemein. (Uniform coding of the phenological development stages of mono- and dicotyledonous plants – extended BBCH scale, general). Nachrichtenblatt des Deutschen P�anzenschutzdienstes, 44(12), 265–70. [In German] Hay, R.K.M., O�er, N.W. (1992). Helianthus tuberosus as an alternative forage crop for cool maritime regions: A  preliminary study of the yield and nutritional quality of shoot tissues from perennial stands. Journal of the Science of Food and Agriculture, 60(2), 213–221. https://doi.org/10.1002/ jsfa.2740600209 Incoll, L.D., Neales, T.F. (1970). �e stem as a temporary sink before tuberization in Helianthus tuberosus L. Journal of Experimental Botany, 21(2), 469–476. https://doi.org/10.1093/jxb/21.2.469 Izsáki, Z., Németh, G.K. (2013). Biomass accumulation and nutrient uptake of Jerusalem artichoke (Helianthus Tuberosus L.). American Journal of Plant Sciences, 4(8), https://doi.org/10.4236/ ajps.2013.48197. 217 Johansson, E., Prade, T., Angelidaki, I., Svensson, S.-E., Newson, W.R., Gunnarsson, I.B., Persson-Hov- malm, H. (2015). Economically viable components from Jerusalem artichoke (Helianthus tuberosus L.) in a biore�nery concept. International Journal of Molecular Sciences, 16(4), 8997–9016. https://doi. org/10.3390/ijms16048997 Kays, S.J. Nottingham, S.F. (2008). Biology and chemistry of Jerusalem artichoke: Helianthus tuberosus L. 1st ed. Boca Raton: CRC Press, pp. 496. Kays, S.J., Kultur, F. (2005). Genetic variation in Jerusalem artichoke (Helianthus tuberosus L.) �owering date and duration. Hortscience, 40(6), 1675–1678. https://doi.org/10.21273/HORTSCI.40.6.1675 Kocsis, L., Kaul, ,H-P. Praznik, W., Liebhard, P. (2007). In�uence of harvest date on shoot and tuber yield ofdi�erent Jerusalem artichoke (Helianthus tuberosus L.) cultivars in the semiarid production area of Austria. Germany Journal of Agronomy, 11, 67–76. Kowalczyk-Juśko, A., Jóźwiakowski, K., Gizińska, M., Zarajczyk, J. (2012). Jerusalem artichoke (Helianthus tuberosus L.) as renewable energy raw material. TEKA. Commission of Motorization and Energetics in Agriculture, 12(2), 117–121. Kulczyński, B., Gramza-Michałowska, A. (2016). Właściwości prozdrowotne fruktanów typu inuliny (Health-promoting properties of inulin-type fructans). Medycyna Rodzinna, 19(2), 86–90. [In Polish] Large, E.C. (1954). Growth stages in cereals illustration of the feekes scale. Plant Pathology, 3(4), 128–129. https://doi.org/10.1111/j.1365-3059.1954.tb00716.x Łukasiewicz, S. (2006). A  modi�cation suggestion of the method of drawing the wet “humid” period in “the Walter’s climate diagram”. Badania �zjogra�czne nad Polską Zachodnią seria A  – geogra�a �zyczna, 57, 95–99. Ma, X.Y., Zhang, L.H., Shao, H.B., Xu, G., Zhang, F., Ni, F.T., Brestic, M. (2011). Jerusalem artichoke (Helianthus tuberosus), a medicinal salt-resistant plant has high adaptability and multiple-use values. Journal of Medicinal Plants Research, 5(8), 1272–1279. Maj, G., Piekarski, W., Słowik, T. (2013). Topinambur (Helianthus tuberosus) substratem do produkcji bi- ogazu (Topinambur (Helianthus tuberosus) as a substrate for biogas production). Gaz, Woda; Technika Sanitarna, 1(2), 59. [In Polish] Mazza, G. (1985). Distribution of sugars, dry matter and protein in Jerusalem artichoke tubers. Canadian Institute of Food Science and Technology Journal, 18(3), 263–265. https://doi.org/10.1016/S0315- 5463(85)71927-X McLaurin, W.J., Sombda, Z.C., Kays, S.J. (1999). Jerusalem artichoke growth, development, and �eld storage. I numerical assessment of plant part development and dry matter acquisition and allocation. Journal of Plant Nutrition, 22(8), 1303–1313. https://doi.org/doi.org/10.1080/01904169909365714 Meier, U. (1985). Die merkblattserie 27 entwicklungsstadien von p�anzen. Biologische Bundesanstalt Für Land– Und Forstwirtscha� (�e lea�et series 27 developmental stages of plants. Federal Biological Institute For Agriculture And Forestry). Nachrichtenblatt des Deutschen P�anzenschutzdienstes, 37, 76–77. [In German] Meier, U. (1997). BBCH-Monograph. Growth Stages of Plants – Entwicklungsstadien von P�anzen – Estadios de Las Plantas – Développement Des Plantes. Blackwell Wissenscha�sverlag, Berlin and Wien, p. 1–622. Meier, U. (2018). Growth stages of mono- and dicotyledonous plants. BBCH monograph. Quedlinburg: Julius Kühn-Institut (JKI). https://doi.org/10.5073/20180906-074619 Meier, U., Bleiholder, H., Buhr, L., Feller, C., Hack, H., Heß, M., Lancashire, P.D., Schnock, U., Stauß, R., van den Boom, T., Weber, E., Zwerger, P. (2009). �e BBCH system to coding the phenological growth stagesof plants – history and publications. Journal Für Kulturp�anzen, 61(2), 41–52. P henological grow th stages and B B C H -identification keys of Jerusalem artichoke (H elianthus tuberosus L.) A ng el ik a K lis zc z 218 Meng, Q., Lu, C., Gao, H., Chen, G., Wu, L., Wu, L., Li, S., He, B.-F. (2021). E�cient biosynthesis of exo- polysaccharide from Jerusalem artichoke using a novel strain of Bacillus velezensis LT-2. Bioresource Technology, 320, 124346. https://doi.org/10.1016/j.biortech.2020.124346 Mystkowska, I., Zarzecka, K. (2013). Wartość odżywcza i prozdrowotna słonecznika bulwiastego (Helian- thus tuberosus L.) (Nutritional value and pro-healthy properties of jerusalem artichoke (Helianthus tuberosus L.)). Postępy Fitoterapii, 23, 123–126. [In Polish] Mystkowska, I., Zarzecka, K., Gugała, M., Baranowska, A. (2015). Właściwości probiotyczne i farmakolo- giczne słonecznika bulwiastego (Helianthus tuberosus L.) (Probiotic and pharmacological properties of Jerusalem artichoke (Helianthus tuberosus L.)). Problemy Higieny i Epidemiologii, 96(1), 64–66. [In Polish] Pacanoski, Z., Mehmeti, A. (2020). �e �rst report of the invasive alien weed Jerusalem artichoke (Helian- thus tuberosus L.) in the Republic of North Macedonia. �e Journal Agriculture and Forestry, 66(1), 115–127. https://doi.org/10.17707/AgricultForest.66.1.12 Paungbut, D., Jogloy, S., Vorasoot, N., Patanothai, A. (2015). Growth and phenology of Jerusalem artichoke (Helianthus tuberosus L.). Pakistan Journal of Botany, 47(6), 2207–2214. Pawłowski, B., Jasiewicz, A. (1971). Flora Polska, (Flora of Poland) tom 12. Warszawa: Państwowe Wy- dawnictwo Naukowe, p. 194–195. [In Polish] Podlaski, S., Pietkiewicz, S., Choluj, D., Horaczek, T., Wisniewski, G., Gozdowski, D., Kalaji, H.M. (2017). �e relationship between the soil water storage and water-use e�ciency of seven energy crops. Pho- tosynthetica, 55(2), 210–218. https://doi.org/10.1007/s11099-017-0697-0 Puangbut, D., Jogloy, S., Vorasoot, N., Srijaranai, S., Kesmala, T., Patanothai, A. (2012). In�uence of planting date and temperature on inulin content in Jerusalem artichoke (Helianthus tuberosus L.). Australian Journal of Crop Science, 6, 1159–1165. Puttha, R., Sanun, J., Bhalang, S., Preeya, P.W., �awan, K., Aran, P. (2013). Variations in morphological and agronomic traits among Jerusalem artichoke (Helianthus tuberosus L.) Accessions. Genetic Resources and Crop Evolution, 60(2), 731–46. https://doi.org/10.1007/s10722-012-9870-2 Radovanovic, A.M., Milovanovic, O.Z., Z., Kipic, M.Z., Ninkovic, M.B., Cupara, S.M. (2014). Character- ization of bread enriched with jerusalem artichoke powder content. Journal of Food and Nutrition Research, 2(12), 895–898. https://doi.org/10.12691/jfnr-2-12-6 Rajan, S., Tiwari, D., Singh, T., Saxena, P., Singh, S., Reddy, N., Upreti, K.K., Burondkar, M., Bhag- wan, A., Kennedy, R. (2011). Application of extended BBCH scale for phenological studies in mango (Mangifera indica L.). Journal of Applied Horticulture, 13, 72–78. https://doi.org/10.37855/ jah.2011.v13i02.25 Rossini, F., Loreti, P., Provenzano, M.E., De Santis, D., Ruggeri, R. (2016). Agronomic Performance and beer quality assessment of twenty hop cultivars grown in central Italy. Italian Journal of Agronomy, 11(3), 180–187. https://doi.org/10.4081/ija.2016.746 Rossini, F., Provenzano, M.E., Kuzmanovic, L., Ruggeri, R. (2019). Jerusalem artichoke (Helianthus tuberosus L.): A versatile and sustainable crop for renewable energy production in Europe. Agronomy, 9(9), 528. https://doi.org/10.3390/agronomy9090528 Ruttanaprasert, R., Banterng, P., Jogloy, S., Vorasoot, N., Kesmala, T., Kanwar, R.S., Holbrookand, C.C., Patanothai, A. (2014).Genotypic variability for tuber yield, biomass, and drought tolerance in Jerusalem artichoke germplasm. Turkish Journal of Agriculture and Forestry, 38, 570–580. Sawicka, B., Skiba, D., Kiełtyka-Dadasiewicz, A., Danilčenko, H. (2019). Jerusalem artichoke (Helianthus tuberosus L.) as energy raw material. Proceedings of the International Scienti�c Conference Rural Development, December, p. 336–342. 219 Sawicka, B., Skiba, D., Kotiuk, E. (2012). Wielokierunkowe wykorzystanie surowców ze Słonecznika bul- wiastego (Helianthus tuberosus L.) (Multidirectional use of raw materials from Jerusalem artichoke (Helianthus tuberosus L.)). In: J. Kondracki, K., Skrzyczyńska, K., Zarzecka (eds.), Współczesne dylematy polskiego rolnictwa, 1st ed., 2:332–43. Biała Podlaska: PWS JPII. [In Polish] Serieys, H., Souyris, I., Gil, A., Poinso, B., Bervillé, A. (2010). Diversity of Jerusalem Artichoke clones (Helianthus tuberosus L.) from the INRA-Montpellier collection. Genetic Resources and Crop Evolution, 57(8), 1207–1215. https://doi.org/10.1007/s10722-010-9560-x. Singh, A.K., Bajpai, A., Rajan, S., Das, S.S., Mishra, K.K. (2021). Modi�ed BBCH codi�cation and correla- tion of phenological characteristics with climatic variables in Jamun (Syzigium cuminii skeels). Scientia Horticulturae, 283, 110081. https://doi.org/10.1016/j.scienta.2021.110081 Skiba, D., Sawicka, B. (2016). �e in�uence of the genetic properties and the content of selected minerals in tubers of Helianthus tuberosus L. Herbalism, 1(2), 18. [In Polish] Spitters, C.J.T., Lootsma, M., van de Waart, M. (1988). �e contrasting growth pattern of early and late vari- eties in Helianthus tuberosus, in topinambour (Jerusalem artichoke). Report EUR 11855. Luxembourg: Commission of the European Communities. Stapor, A. (2020). Uprawa topinamburu, czyli zdrowie i piękno z natury sposobem na rozwój obszarów wiejskich. Krajowa Sieć Obszarów Wiejskich. 12 October 2020. http://swietokrzyskie.ksow.pl/aktu- alnosc/uprawa-topinamburu-czyli-zdrowie-i-piekno-z-natury-sposobem-na-rozwoj-obszarow-wiej- skich. [In Polish] Swanton, C.J., Cavers, P.B. (1989). Biomass and nutrient allocation patterns in Jerusalem artichoke (He- lianthus tuberosus). Canadian Journal of Botany, 67(10), 2880–2887. https://doi.org/10.1139/b89-369 Swanton, C.J., Clements, D.R., Moore, M.J., Cavers, P.B. (1992). �e biology of Canadian weeds. 101. Helianthus tuberosus L. Canadian Journal of Plant Science, 72(4), 1367–1382. https://doi.org/10.4141/ cjps92-169 Terzić, S., Altagić, J., Maksimović, I., Zeremski, T., Petrović, S., Dedić, B. (2012) In�uence of photoperiod on vegetation phases and tuber development in topinambour (Helianthus tuberosus L.). Archives of Biological Sciences, 64(1), 175-182. https://doi.org/10.2298/ABS1201175A Tokarska-Guzik, B., Dajdok, Z., Zając, M., Zając, A., Urbisz, A., Danielewicz, W., Hołdyński, C. (2012). Rośliny obcego pochodzenia w Polsce ze szczególnym uwzględnieniem gatunków inwazyjnych (Plants of foreign origin in Poland, with particular emphasis on invasive species). Warszawa: �e General Direc- torate for Environmental Protection. [In Polish] Treder, W., Klamkowski, K., Wójcik, K. (2018). A  new approach to the method of drawing the gauss- en-walter climate diagram. Meteorology Hydrology and Water Management, 6(2), 1–7. https://doi. org/10.26491/mhwm/82237 Troitzky, N.N. (1925). Vorläu�ge Untersuchungsmittel der experimentell-biologischen Station für ange- wandte Entomologie (Preliminary research means of the experimental biological station for applied entomology). P�anzenschutz-Nachrichten Bayer, 32, 97–163. [In German] van Wyk, B.-E., Wink, M. (2008). Rośliny lecznicze świata (Medicinal plants of the world). Wrocław: Med- Pharm, p. 412. [In Polish] Vaughan, J.G., Geissler, C.A. (2001). Rośliny jadalne (Edible plants). Warszawa: Prószyński i S-ka, p. 188. [In Polish] Walter, H. (1976). Strefy roślinności a klimat (Vegetation zones and climate). Warszawa: PWRiL. [In Polish] Westley, L.C. (1993). �e e�ect of in�orescence bud removal on tuber production in Helianthus tuberosus L. (Asteraceae). Ecology, 74(7), 2136–2144. https://doi.org/10.2307/1940858 Zadoks, J.C., Chang, T.T., Konzak, C.F. (1974). A  decimal code for the growth stages of cereals. Weed Research, 14(6), 415–421. https://doi.org/10.1111/j.1365-3180.1974.tb01084.x P henological grow th stages and B B C H -identification keys of Jerusalem artichoke (H elianthus tuberosus L.) A ng el ik a K lis zc z 220 Zhang, A., Zang, W., Zhang, X., Ma, Y., Yan, X., Pang, Q. (2016). Global proteomic mapping of alkali stress regulated molecular networks in Helianthus tuberosus L. Plant and Soil, 409(1), 175–202. https://doi. org/10.1007/s11104-016-2945-7 Zhao, G., Gao, Y., Gao, S., Xu, Y., Liu, J., Sun, C., Gao, Y., Liu, S., Chen, Z., Jia, L. (2019). Phenological growth stages of Sapindus mukorossi according to BBCH scale. Forests, 10(462), 1–15. https://doi. org/10.3390/f10060462 Zubr, J. (1988). Jerusalem artichoke as a �eld crop in Northern Europe, in topinambour (Jerusalem artichoke). Report EUR 11855. Luxembourg: Commission of the European Communities. 221 Appendix 1 Tab. 2. Phenological growth stages of Helianthus tuberosus L. according to the BBCH scale BBCH code (2-digit) Description of the development from the tuber Principal growth stage 0: Sprouting 00 Innate or enforced dormancy, tuber not sprouted 01 Beginning of sprouting: �rst sprout visible (< 1 mm) 02 End of dormancy: sprout 2–3 mm 03 First (main) sprout further growth (< 1 cm) 04 Beginning of root formation; �rst sprout further growth (< 5 cm) 05 First sprout further growth (< 15 cm) 06 First sprout further growth (< 15 cm) 07 First sprout further growth (< 20 cm) 08 First sprout further growth (< 30 cm and more) 09 Breakthrough sprouts on the ground Principal growth stage 1: Leaf development (main shoot) 10 �e �rst pair of leaves begins photosynthesis (leaves are still rolled up, but green) 11 First pair of leaves fully developed 12 Second pair of leaves fully developed 13 �ird pair of leaves fully developed 14 Fi�h pair of leaves fully developed 15 Seventh pair of leaves fully developed 16 Ninth pair of leaves fully developed 17 Eleventh pair of leaves fully developed 18 �irteenth pair of leaves fully developed 19 Fi�eenth pair of leaves fully developed 19n Sixteenth (and further) pair of leaves fully developed Principal growth stage 2: Formation of other sprouts 20 Only the main shoot is visible above the ground 21 �e �rst side shoot appears above the ground 22 �e second side shoot appears above the ground 2.. �e next side shoots appear above the ground Principal growth stage 3: Development of lateral branches on main shoot (Side branching) 30 Beginning of developing lateral branches: the �rst (twin) axillary buds appear in the axils of the �rst pair of leaves on the main shoot 31-1 �e �rst pair of developed leaves growing out from the axils of the �rst pair of leaves on the main shoot 31-2 �e second pair of developed leaves growing out from the axils of the �rst pair of leaves on the main shoot 31-… Successive pairs of developed leaves growing out from the axils of the �rst pair of leaves on the main shoot P henological grow th stages and B B C H -identification keys of Jerusalem artichoke (H elianthus tuberosus L.) A ng el ik a K lis zc z 222 32-0 �e �rst (twin) axillary buds appear in the axils of the second pair of leaves on the main shoot 32-1 �e �rst pair of developed leaves growing out from the axils of the second pair of leaves on the main shoot 32-.. Successive pairs of developed leaves growing out from the axils of the second pair of leaves on the main shoot 33-0 �e �rst (twin) axillary buds appear in the axils of the third pair of leaves on the main shoot 33-1 �e �rst pair of developed leaves growing out from the axils of the third pair of leaves on the main shoot 33-.. Successive pairs of developed leaves growing out from the axils of the third pair of leaves on the main shoot 34-0 �e �rst (twin) axillary buds appear in the axils of the ��h pair of leaves on the main shoot 34-1 �e �rst pair of developed leaves growing out from the axils of the ��h pair of leaves on the main shoot 34-.. Successive pairs of developed leaves growing out from the axils of the ��h pair of leaves on the main shoot 35-0 �e �rst (twin) axillary buds appear in the axils of the seventh pair of leaves on the main shoot 35-1 �e �rst pair of developed leaves growing out from the axils of the seventh pair of leaves on the main shoot 35-.. Successive pairs of developed leaves growing out from the axils of the seventh pair of leaves on the main shoot 36-0 �e �rst (twin) axillary buds appear in the axils of the ninth pair of leaves on the main shoot 36-1 �e �rst pair of developed leaves growing out from the axils of the ninth pair of leaves on the main shoot 36-.. Successive pairs of developed leaves growing out from the axils of the ninth pair of leaves on the main shoot 37-0 �e �rst (twin) axillary buds appear in the axils of the eleventh pair of leaves on the main shoot 37-1 �e �rst pair of developed leaves growing out from the axils of the eleventh pair of leaves on the main shoot 37-.. Successive pairs of developed leaves growing out from the axils of the eleventh pair of leaves on the main shoot 38-0 �e �rst (twin) axillary buds appear in the axils of the thirteenth pair of leaves on the main shoot 38-1 �e �rst pair of developed leaves growing out from the axils of the thirteenth pair of leaves on the main shoot 38-.. Successive pairs of developed leaves growing out from the axils of the thirteenth pair of leaves on the main shoot 39-0 �e �rst (twin) axillary buds appear in the axils of the ��eenth pair of leaves on the main shoot 39-1 �e �rst pair of developed leaves growing out from the axils of the ��eenth pair of leaves on the main shoot 39-.. Successive pairs of developed leaves growing out from the axils of the ��eenth pair of leaves on the main shoot 39n-0 �e �rst (twin) axillary buds appear in the axils of the sixteenth (or successive) pair of leaves on the main shoot 223 Principal growth stage 4: Rhizomes booting 40 �e �rst rhizome starts to grow 41 �e other rhizomes start to develop 42 �e rhizomes elongate 43 �e rhizomes still elongate and start to branch 44 �e rhizomes elongate and branch still (< 30 cm from plant) 45 �e rhizomes elongate and branch still (< 50 cm from plant) 46 �e rhizomes elongate and branch still (< 80 cm from plant) 47 �e rhizomes elongate and branch still (< 100 cm from plant) 48 Rhizomes are developed and most of them have not thickened ends yet 49 �e ends of most rhizomes start to thicken (rhizome tips to twice the diameter of subtending rhizome) Principal growth stage 5: In�orescence emergence 50 In�orescence not visible 51 In�orescence just visible between youngest leaves on the main shoot (tight bud stage) (peduncles elongate) In�orescence just visible on the upper branches (50% branches on the top have in�o- rescence just visible); tight bud stage (peduncles elongate) In�orescence just visible on the lower branches; tight bud stage (peduncles elongate) 52 First top ligules (ray �ower corollas) exposed and green (peduncles elongate) 53 Most top ligules exposed and green (peduncles elongate) 54 Ligules on the upper branches exposed and green (peduncles elongate) 55 Most top ligules exposed and green (peduncles elongate) 56 Ligules on the upper branches exposed and green (peduncles elongate) 57 Ligules on the lower branches exposed and green (peduncles elongate) 58 Ligules on the top and upper branches yellow-green (in�orescence still closed) 59 Most ligules on the plant are yellow-green (in�orescence still closed) Principal growth stage 6: Floweringi 60 Top ligules beginning to unroll (disk �ower corollas yellow and closed) 61 First top ligules open 62 First top ligules with emerging anthers from the corolla 63 Additional anthers and �rst stigmas emerging on outer whorls on the top ligules 64 About half of top disk �owers open with stigmas emerged 65 All of the top disk �owers open with stigmas emerged (in bloom) 66 �ird part of disk �owers in lateral branches (from the top) are in bloom (outer whorl �owers on the top displaying initial stigma senescence) 67 Two thirds of disk �owers on lateral branches (from the top) are in bloom (top ligules wilting and initial drying) 68 80% of disk �owers on lateral branches (from the top) are in bloom 69 End of �owering: almost all disc �ower have �nished �owering on the plant, ray �orets dried Principal growth stage 7: Tubers bulking 70 Tubers bulking (10% of all) 71 Tubers bulking (20% of all) 72 Tubers bulking (30% of all) 73 Tubers bulking (40% of all) P henological grow th stages and B B C H -identification keys of Jerusalem artichoke (H elianthus tuberosus L.) A ng el ik a K lis zc z 224 74 Tubers bulking (50% of all) 75 Tubers bulking (60% of all) 76 Tubers bulking (70% of all) 77 Tubers bulking (80% of all) 78 Tubers bulking (90% of all) 79 Tubers are full (100%), maximum of the total tuber mass reached Principal growth stage 8: Ripening seedii Principal growth stage 9: Senescence 90 40% of aboveground green parts of the plant has dried up 91 50% of aboveground green parts of the plant has dried up 92 60% of aboveground green parts of the plant has dried up 93 70% of aboveground green parts of the plant has dried up 94 80% of aboveground green parts of the plant has dried up 95 90% of aboveground green parts of the plant has dried up 96 100% of aboveground green parts of the plant has dried up 97 Aboveground biomass has > 20% moisture w/w 98 Aboveground biomass has < 20% moisture w/w 99 Tuber harvested, dormancy i a synthesis of an in�orescence development a�er Kays and Nottingham (2008) (changed) ii seed is developing along the tuber bulking 225 Fazy rozwojowe słonecznika bulwiastego (Helianthus tuberosus L.) w propozycji oznaczeń skali BBCH Streszczenie Celem pracy było zbadanie i standaryzacja faz rozwojowych słonecznika bulwiastego (Helianthus tuberosus L.), rosnącego w klimacie umiarkowanym, na podstawie klucza oznaczeń BBCH. Tego rodzaju analizę wykonano po raz pierwszy, co było oczekiwane w dyskursie naukowym, jak i praktycznym. Rosnące zainteresowanie tym gatunkiem, zarówno z punktu widzenia naukowego, jak i utylitarnego, stawia potrzebę nazwania jego poszczególnych stadiów rozwojowych oraz ich standaryzacji w zakresie nomenklatury. Słonecznik bulwiasty jest rośliną o wielokierunkowym wykorzystaniu w różnych gałęziach przemysłu. Surowcem w przemyśle spożywczym są bulwy, które gromadzą znaczne ilości inuliny – łańcuchowego polimeru fruktozy, o istot- nych właściwościach probiotycznych. Dzięki temu bulwy są cennym składnikiem żywności funkcjonalnej, substratem w produkcji farmaceutyków, czy napojów alkoholowych, a także pozwalają na przetrwanie gatunku w środowisku w okresie zimowym. Formowanie się bulw zachodzi przez znaczną część rozwoju ontologicznego gatunku (BBCH 49) i związane jest głównie z fotoperiodem, sumą temperatur efektywnych oraz obecnością nadziemnej biomasy rośliny, z której zachodzi alokacja asymilatów do bulw w okresie rozwoju generatywnego. Spośród wielu innych zastosowań, roślina ta jest wykorzystywana jako surowiec energetyczny, gdyż naturalnie wyschnięta biomasa nadziemna, pod koniec sezonu wegetacyjnego, zawiera niską zawartość wody i plasuje ten gatunek w środku listy roślin energetycznych. Key words: topinambour, BBCH scale, phenological stages Received: [2021.06.02] Accepted: [2021.09.17] P henological grow th stages and B B C H -identification keys of Jerusalem artichoke (H elianthus tuberosus L.)