ISSN: 2312-3370, Agricultural Science and Practice, 2015, Vol. 2, No. 1

UDC 582.4 : 582.662 + 581.9 + 581.522.6 + 581.527 AS A PROMISING PSEUDOCEREAL CROP FOR UKRAINE S. L. Mosyakin, V. V. Schwartau 1 M. G. Kholodny Institute of Botany, National Academy of Sciences of Ukraine 2, Tereshchenkivska Str., Kyiv, Ukraine, 01601 2 Institute of Plant Physiology and Genetics, National Academy of Sciences of Ukraine 31/17, Vasylkivska Str., Kyiv, Ukraine, 03022 e-mail: [email protected]; [email protected] Received on February 09, 2015

The article provides an assessment of perspectives of quinoa (Chenopodium quinoa L.) cultivation in Ukraine, based on international experience and original ¿ eld tests, with the aim of ensuring further development and diversi¿ cation of crop production in Ukraine and expanding modern crop rotation systems. The data on the taxonomic position of quinoa and its relationships with other species of the genus Chenopodium and the history of species are provided. Quinoa is a crop of high nutritional value and can be used in gluten-free diets, which are important components of human ration. The results of test cultivation of quinoa in 2013–2014 under conditions of the experimental agricultural farm of the Institute of Plant Physiology and Genetics, Na- tional Academy of Sciences of Ukraine, located in Vasylkiv District (rayon) of Kyiv Region (oblast), are pro- vided. It is concluded that quinoa is a promising crop for domestic grain producers. The introduction of quinoa into crop rotation systems can improve ecological conditions of agroecosystems and promote restoration of soil fertility in the country without diminishing the revenues of farmers. Key words: Chenopodium quinoa, quinoa, biology, taxonomy, domestication, agricultural technologies.

INTRODUCTION Except for the vegetation season of 2014, during the recent ¿ ve years Ukraine witnessed numerous long- The Big Three – , and corn – are term drought periods, which sharply decreased crop justly considered to be the main crops and feed- yield of spiked cereals, technical and other crops. Each ers of humanity. According to FAO (http://www.fao. year the ¿ elds of the Steppe zone of Ukraine suffer org), cereals account for about 58 % of the annual crop from droughts of varied intensity. Rapid changes in the areas and provide humans with over 50 % of food calo- structure of Ukrainian caused by the forma- ries. By 2050, the share of the three mentioned crops tion of agroholdings (agricultural holding companies together is expected to amount up to 80 % of the in- and corporations) and the decline of animal farming crease in cereal consumption [1, 2]. However, it is ac- resulted in reduced crop rotations with the cultivation knowledged that along with the increase in the cultiva- of only immediately pro¿ table and cost-ef¿ cient crops tion of the main crops there is a need for diversi¿ cation with high nutrient removal from soils. This, in turn, re- in cultivation and consumption of other crops, includ- sulted in considerable worsening of phytosanitary con- ing those currently of lesser signi¿ cance (so-called mi- ditions of agroecosystems and segetal plant communi- nor crops), but which were in the past, and still are, ties, the increase in weed infestation levels and growing components of traditional agricultural systems [2–5]. threats of emergence of pesticide-resistant pathogens, These crops may be better adjusted to the conditions pests, and weeds, as well as the decline of soil fertility. of speci¿ c geographic regions of the Globe, where the A considerable share of ¿ elds is occupied cultivation of the main crops is complicated, risky, or by cultivars and hybrids of foreign selection, which economically or ecologically unreasonable. often become frostbitten in vast areas. Among staple

AGRICULTURAL SCIENCE AND PRACTICE Vol. 2 No. 1 2015 3 MOSYAKIN et al. cereals registered in Ukraine, spring wheat does not The taxonomic position of quinoa and match the main domestic crop – winter wheat – in the its relationships to other species of Chenopodium yield level and qualitative indices, while other crops According to modern views, the family Chenopodia- (peas, for instance) also require the application of great ceae Vent. belongs to the order Caryophyllales Juss. ex amounts of pesticides. Bercht. & J. Presl of the unranked group of eudicot, or It is important to amend crop rotations in Ukraine true dicotyledonous angiosperms [8, 9]. According to with highly cost-ef¿ cient spring crops with low level our variant of the system of À owering plants [9], the of nutrient removal, which are resistant to high tem- taxonomic position of Chenopodiaceae is viewed as peratures and droughts and tolerant to the applica- follows: division Magnoliophyta Cronquist, Takht. & tion of agricultural chemicals. Therefore, so-called W. Zimmerm. ex Reveal (angiosperms, or À owering pseudocereals nowadays appear to be quite promising plants), class Rosopsida Batsch (= Dicotyledonae, eu- food and technical crops. Pseudocereals are de¿ ned dicots), subclass Caryophyllidae Takht., order Caryo- as non-grass crops that can be used in much the same phyllales Juss. ex Bercht. & J. Presl., and suborder way as true cereals (grass cereals). It means that their Chenopodiineae J. Presl. seeds (“grain”) are used similarly to cereal grain, e.g., In all three published variants of the APG system, the pseudocereal seeds can be ground into À our, grits, family Chenopodiaceae [8, 10, 11] was merged with etc. Typical pseudocereals include (Fago- the family Amaranthaceae, the latter name having no- pyrum esculentum Moench, family Polygonaceae), menclatural priority and thus being used for the com- grain (several domesticated species of the bined family. However, it is dif¿ cult to agree with this genus Amaranthus L., family Amaranthaceae), as decision, especially considering available taxonomic well as quinoa (Chenopodium quinoa L.) and some and nomenclatural evidence [9, 12]. More detailed other species of the genus Chenopodium L. (family justi¿ cation of the need to preserve the independence Chenopodiaceae). of the family Chenopodiaceae will be published in an- Quinoa, an ancient crop that emerged approximately other article (Mosyakin, in print). 7,000 years BP (before present) in the mountain re- Within the family, the genus Chenopodium be- gions of the central part of the Andes in South America, longs to subfamily Chenopodioideae Burnett, tribe currently enjoys the period of its revival and renewed Chenopodieae Dumort. It has been suggested to popularity. It is caused by unique food (nutrient) char- synonymize the latter with tribe Atripliceae Duby acteristics of this species, its easiness of cultivation, [13], but it seems to be not the best option (Mosya- environmental resistance and tolerance, and a great kin, in print). number of available and diverse cultivars [2, 3, 5, 6]. The modern system of Chenopodium sensu lato and This crop was even used as an experimental object in other genera previously included into this aggregate is the NASA space program [7]. Therefore, the United based on the system of Aellen, a Swiss botanist [14– Nations General Assembly and FAO declared 2013 as 16]; that system was later improved and updated by the International Year of Quinoa (http://www.fao.org/ Scott [17], further on – by one of the authors of this quinoa-2013/en/). article [18–21], in particular, in collaboration with the As for nutritional properties of quinoa, its protein US botanist Clemants [22, 23]. Mosyakin and Clem- content in “grain” varies from 13.81 to 21.9 %, depend- ants [22, 23], among other taxonomic and nomencla- ing on a cultivar. Quinoa is one of a few edible plants tural proposals, ¿ rst justi¿ ed the phylogenetic isolation containing all essential amino acids and most closely of so-called glandular chenopods (aromatic species corresponding to human food standards approved by with glandular trichomes) from typical representatives FAO. The balance of essential amino acids in quinoa of Chenopodium sensu stricto (in the strict sense) and protein considerably exceeds the indices of amino acid transferred these species to a re-circumscribed genus content in wheat grain, , and soybean, approach- Dysphania R. Br., providing necessary nomenclatural ing in that respect the parameters of milk protein. Qui- combinations. This taxonomic decision was initially noa also has signi¿ cantly higher calcium, magnesium, accepted in our À oristic and taxonomic treatments for iron and zinc contents as compared to those values in the Flora of North America North of Mexico and the wheat, corn, rice, barley, , , and , among Flora of China [24, 25], later con¿ rmed by molecular others. It is important that quinoa is suitable for gluten- phylogenetic data [13, 26í28], and currently it is al- free diet [5]. most universally accepted.

4 AGRICULTURAL SCIENCE AND PRACTICE Vol. 2 No. 1 2015 QUINOA AS A PROMISING PSEUDOCEREAL CROP FOR UKRAINE In addition, modern molecular phylogenetic studies In pre-Columbian times (i.e. prior to the beginning justi¿ ed further segregation of several additional gen- of colonization by the Europeans) ancient inhabitants era from Chenopodium sensu lato (in the wide sense) of America managed to domesticate quite a wide range [13, 28, 29]. However, these radical changes do not af- of local wild plants; however, these crops considerably fect the group that includes quinoa and other pseudo- differed taxonomically from the domesticated plants of cereals of the genus, since they all remain in Chenopo- Eurasia. The unique feature of the ancient American dium in the strict sense. domestication experience is the use of exclusively local According to modern taxonomic views [30, 31], qui- resources, especially under very limited contacts even noa belongs to sect. Chenopodium (the typical section, between American geographical and cultural centers of containing the nomenclatural type of the genus), which domestication. Several main centers of plant domes- includes most species of the genus further segregated tication and origin of agriculture, with their peculiar into several subsections. Subsection Favosa (Aellen) sets of crops, are distinguished in the Americas (North, Mosyakin & Clemants (= Chenopodium series Fa- Central, and South) (see reviews for further informa- vosa Aellen) includes species geographically ranging tion [45–53]). from South America (C. quinoa Willd., C. hircinum The following centers of plant domestication are usu- Schrad. etc.) to North America (a group of species re- ally distinguished, with some variants, in the Americas: lated to C. berlandieri Moq.) and Eastern and South ƒ Mesoamerican upland and Mesoamerican lowland Asia (C. ¿ cifolium Sm. sensu lato). The latter species centers (remarkable domesticates are corn Zea mays L., (the lectotype of subsect. Favosa) was widely natural- grain L. = A. panicula- ized in other regions outside its initial natural range, tus L., species of such genera as beans Phaseolus, squash due to which it was sometimes erroneously considered and pumpkin Cucurbita, paprika Capsicum etc.); a species of Mediterranean origin. The phylogenetic integrity of the newly outlined subsect. Favosa was ƒ Andean center (potato Solanum tuberosum L. s.l. conclusively supported by modern molecular and phy- (some taxa), ulluco Ullucus tuberosus Caldas, oca Ox- logenetic evidence [13, 28] and comparative anatomic alis tuberosa Molina; quinoa Chenopodium quinoa and carpological data [32]. Willd., qañiwa C. pallidicaule Aellen, foxtail amaranth Amaranthus caudatus L. etc.); Issues of taxonomy and evolution of several species of Chenopodium, in particular, American cultivated ƒ Arizonian-Sonoran center (formed under the inÀ u- races of quinoa (C. quinoa), huazontle (C. nuttalliae ence of the Mesoamerican upland center; also includes Safford), and their wild relatives (C. hircinum etc.) speci¿ c cereals at early stages of domestication, in par- and probable ancestors were considered in some sub- ticular, barnyard from the Echinochloa muricata stantial, as of that time, publications by Wilson and his (P. Beauv.) Fern. group, species of millet Panicum s.l., colleagues [35í42]. Even at that time, these integral barley Hordeum s.l., bromegrass Bromus s.l. etc.); studies involved the methods of traditional taxonomy, ƒ Eastern North American, also known as Alaba- palaeoethnobotany (archaeobotany), biochemistry, and man-Illinoian, center ( Moq. – biosystematics. C. nuttallii aggr., species of genera Helianthus, Cy- Recently the nomenclatural type (lectotype) of the clachaena, Ambrosia, Polygonum s.l., Phalaris etc.); species has been selected for C. quinoa [43]; this her- ƒ Peruvian coastal center (mainly adoption of crops barium specimen (the reference sample of the species) from the Andes); is deposited at the Willdenow Herbarium of the Botani- cal Garden and Museum in Berlin (Botanischer Garten ƒAmazonian, also known as Amazonian-Orinocan or und Botanisches Museum Berlin-Dahlem). Brazilian-Paraguayan, center (manioc Manihot escu- lenta Crantz, sweet potato Ipomoea batatas (L.) Lam., History and peculiarities of domestication and some other crops). of quinoa and some other unique crops of America As we can see, the absolute majority of domesticates The issues of history and peculiarities of domestication of American origin, especially those used at the stage of quinoa and some other unique crops of America has of early agriculture, are remarkable for their ruderal already been considered in detail by one of the authors (explerent) life strategy; by their ecological and phy- of this article [44]; subsequent discussion in the present tosociological features they are somewhat similar to article is partially based on that earlier publication. weeds or plants of marginal habitats [44].

AGRICULTURAL SCIENCE AND PRACTICE Vol. 2 No. 1 2015 5 MOSYAKIN et al. Species of families Chenopodiaceae and Amarantha- jonesianum B.D. Smith (the subspecies considered al- ceae occupy a prominent place among the cultivated ready extinct), and some others. Among the mentioned native plants of America [6, 35, 36, 39í42, 49, 52í59]. species and races, only quinoa still maintains its eco- The independent domestication of ruderal species of nomic importance, while remaining species have been Chenopodiaceae and Amaranthaceae in America can either forgotten (for instance, domesticated North be viewed as a unique domestication experiment. The American large-seeded races of the C. berlandieri documented role of these domesticated plants in Ame- group), or are preserved only locally, as relict crops. rica was much more important than the role of crops The majority of domesticated species and subspe- of these families in the Old World. With the exception cies of Chenopodium demonstrate some morphologi- of beet (Beta vulgaris L.) and spinach (Spinacia ole- cal and physiological changes likely caused by do- racea L.) that originated in the Mediterranean region mestication: disruption of the normal mechanism of [60, 61], chenopod crops did not have signi¿ cant distri- spontaneous dissemination (however, seeds are easily bution in the Old World, mainly being used there from shed while being thrashed), lost (or at least decreased) time to time as leaf crops (salad plants) or as substitutes dormant period in seeds; the sizes of seeds increase of normal and wholesome food in famine years (so- considerably; the pericarp is easily isolated from the called famine food). testa (seed coat), the testa itself becomes thinner, due The ethnobotanical data on the use of wild repre- to which the “seeds” acquire shades of white, yellow sentatives of Chenopodiaceae and Amaranthaceae by or reddish color. native peoples of North and South America are pres- As we can see, practically all cultivated species of ent in many publications (see above), and similar data Chenopodium were domesticated independently in on North America were critically analyzed and sum- various regions of America. In particular, the fact of in- marized in a thorough monograph by Moerman [63]. dependent origin of Andean cultivars of C. quinoa and Fresh green parts of many representatives of Cheno- Mexican cultivars of C. nuttaliae [42] has been proven podiaceae (species of genera Atriplex and Chenopo- without doubt. Despite numerous studies, the origin of dium) were used as green (salad) vegetables or pot- quinoa is still a matter of dispute. This species is likely herbs. The seeds of many wild species of Cheno- to have originated from one of the local Andean tet- podiaceae served as substitutes of cereals: grit or raploid wild species belonging to subsect. Favosa. At À our were made of them; they were soaked, ground, present the researchers [64, 65] are inclined to think boiled and prepared as porridge, substitute, that tetraploid American species of Chenopodium from and pemmican-type dry mixtures. In particular, Mo- subsect. Favosa (including the cultivated ones) are al- erman [63] described the use of fruits and seeds of lopolyploids that originated from ancient hybridization at least 9 species of Atriplex (orache), 10 species of events between the local American and probably some Chenopodium, species of Cycloloma Moq. (winged Asian (or Eurasian) species. In particular, it is believed pigweed), Corispermum L. (bugseed) etc. as food that one diploid (2n = 18) ancestor of species of the products, but the real number of species actually tetraploid (2n = 36) American complex (C. berlandieri, used must have been much higher, as ethnobotanists C. hircinum, etc.) could be the North American species often did not indicate the precise species-level iden- C. standleyanum Aellen. Eurasian species C. album L. ti¿ cation of plants, reporting only the genus or an- or C. ¿ cifolium were suggested for the role of the sec- other supraspeci¿ c group. ond probable diploid ancestor. In our opinion, typical Archaeological and palaeoethnobotanical data (see C. album, being hexaploid (2n = 54), cannot be consid- references above) indicate wide use of seeds of several ered as a potential parental species of a tetraploid tax- species of Chenopodium, which may be considered true on. Therefore, diploid C. ¿ cifolium is best suited for the domesticates, in North and South America: C. quinoa role of such an ancestral species. The species C. stand- (quinoa, the Andes), C. pallidicaule (qañiwa or qaña- leyanum was placed by us [24, 30] in a separate sub- wa, the Andes), C. nuttaliae Safford (= C. berlandieri section Standleyana Mosyakin & Clemants, which is subsp. nuttalliae (Safford) H.D. Wilson & C.B. Heiser; extremely interesting in terms of its phytogeography, huazontle, Mexico), C. berlandieri (central and south- since it links North American and East Asian taxa. This western part of South America), C. bushianum Aellen subsection also includes East Asian C. bryoniifolium (= C. berlandieri subsp. bushianum (Aellen) Cronq., Bunge (= C. koraiense Nakai s. str.), C. gracilispicum North American Atlantic region), C. bushianum subsp. H.W. Kung (C. koraiense auct. p.p.), C. atripliciforme

6 AGRICULTURAL SCIENCE AND PRACTICE Vol. 2 No. 1 2015 QUINOA AS A PROMISING PSEUDOCEREAL CROP FOR UKRAINE Murr s. str. (C. badachschanicum auct., non Tzvelev), (2) Salares (southern highlands), (3) Inter Andean Val- North American C. standleyanum (C. boscianum auct. leys, (4) arid zones (in particular, western highlands), non Moq.), and probably also C. missouriense Ael- plants of (5) high altitudes and cool climate, (6) coastal len emend. F. DvoĜák. Thus, there is an evident “East regions, (7) jungle and tropical zones, and (8) high Asian trace” in the origin of American tetraploid spe- rainfall and humidity zones. Such a considerable diver- cies of subsect. Favosa (including C. berlandieri, C. sity of cultivars and forms opens excellent prospects hircinum, and C. quinoa). for the search for cultivars best pre-adapted to the con- ditions of different physiographic and climatic zones As currently generally estimated, quinoa emerged of Ukraine. about 7,000 years BP, but there are some indications to an older age of this crop. The archaeobotanical remains Peculiarities of quinoa cultivation of the fruits similar to those of quinoa in the northern part of Peru on the western slopes of the Andes are dat- The culture of quinoa is new for domestic pro- ed by the Las Pircas period (9,800–7,800 years B.P.); ducers in Ukraine. Under the conditions of south- however, it is possible that these earliest ¿ nds indicate ern Polissia and northern Forest-Steppe zones of the pre-domestication use of plants from natural habi- Ukraine – at the experimental agricultural farm of tats by tribes of hunters-gatherers rather than complete the Institute of Plant Physiology and Genetics, NAS domestication [43]. of Ukraine (Vasylkiv District, Kyiv Region) – qui- noa was sown in well prepared leveled soil in early Contrary to the older Andean complex, the so- May in 2013–2014 at stable positive temperatures called “Eastern complex” of crops was formed in (8í10 °ɋ) and suf¿ cient humidity of soil. The row North America in the eastern and central parts of the planting was made 1–2 cm deep with the row spac- present-day USA about 5,000 years ago, i.e. much later than the time of quinoa domestication in South ing of 30 cm. The seeds were kindly provided by Dr. America. Besides the races of the C. berlandieri – C. Jamal B. Rakhmetov (M.M. Gryshko National Bo- bushianum group, this complex also included Am- tanical Garden, NAS of Ukraine). Late terms of sow- brosia tri¿ da L. (giant ragweed), Helianthus annuus ing allow using non-selective herbicides based on N- L. s.l. (sunÀ owers) and local species of genera Cy- (phosphonomethyl)glycine (Glyphosate), in spring clachaena (marsh-elder) and Amaranthus. Consider- and relieving the ¿ elds from wheatgrass, Elytrigia able domestication-caused morphological changes repens (L.) Desv. ex Nevski (Agropyrum repens L.), in the seeds of cultivated species of Chenopodium sow-thistle species (Sonchus spp.), and other weeds. are reliably traced in the archaeological records of Being the predecessor, quinoa allows decreasing the this region for the period of 3,500–2,000 years B.P. weed infestation of the ¿ eld with thermophylic grass During that period, forms with large (about 2 mm species (in particular, foxtail species, Setaria spp. in diameter) seeds and thinner testa (10–15 —m as etc.) on condition of introducing graminicides of the compared to 40–78 —m in wild species) appeared, classes of aryloxyphenoxypropionic acid, cyclohex- which may testify to the emergence of true domes- anediones etc. A considerable drawback is a high ticated cultivars. The cultivation of local Chenopo- level of infestation of quinoa ¿ elds with common dium domesticates (as well as many other species) goosefoot (Chenopodium album L., a species very in North America gradually ceased starting with the widespread in Ukraine) and other related species of 2nd–3rd centuries C.E. due to the adaptation of more the genus, as well as species of the genus Atriplex L. productive Mesoamerican (Central American) crops, (orache). However, if quinoa is used as a predecessor in particular, corn, several species of beans, squash of winter wheat or corn, on the ¿ elds of which the and pumpkin [53]. However, the ancient tradition of use of herbicides to control Chenopodium album is cultivation of South American species of Chenopo- ef¿ cient, high levels of weed biomass accumulation dium (especially quinoa) continued and still contin- may be considered as application of green manure. ues until now. Quinoa responds very well to soil enrichment with Quinoa has many native landraces and other cultivars nitrogen [66]. To cultivate the organic crop, quinoa adapted to extremely diverse natural and climatic con- should be sown after the application of organic fertil- ditions of South America. In particular, the following izers, green manure, nitrogen-¿ xing grain legumes, or groups of cultivars, landraces, and forms are distin- either nitrogen-¿ xing species of plants should be used guished [6]: (1) Altiplano (northern Andean highlands), as green manure.

AGRICULTURAL SCIENCE AND PRACTICE Vol. 2 No. 1 2015 7 MOSYAKIN et al. Quinoa demonstrates good response to the foliar ap- mal for the regions in the Steppe, Forest-Steppe, and plication of macro- and microelements. However, the Polissia zones, the control of dicotyledonous species damage of quinoa leaves with agrochemicals may be of weeds in the ¿ elds, and some aspects of introduc- observed as early as after the introduction of 1–3 % ing the mechanical aids into crop production. working solutions of fertilizers. Foliar bioforti¿ cation Ʉɿɧɨɚ – ɩɟɪɫɩɟɤɬɢɜɧɚ ɞɥɹ ɍɤɪɚʀɧɢ of quinoa yield is ef¿ cient using fertilizers containing «ɡɟɪɧɨɜɚ» ɤɭɥɶɬɭɪɚ nutrition elements and amino acids (Megafol, Megafol ɋ. Ʌ. Ɇɨɫɹɤɿɧ 1, ȼ. ȼ. ɒɜɚɪɬɚɭ 2 Protein, Isabion). e-mail: [email protected]; Due to the fact that quinoa seeds have high content e-mail: [email protected] of proteins and amino acids, it was demonstrated that 1 it may condition the increase in selenium accumulation ȱɧɫɬɢɬɭɬ ɛɨɬɚɧɿɤɢ ɿɦ. Ɇ. Ƚ. ɏɨɥɨɞɧɨɝɨ ɇȺɇ ɍɤɪɚʀɧɢ ȼɭɥ. Ɍɟɪɟɳɟɧɤɿɜɫɶɤɚ, 4, Ʉɢʀɜ, ɍɤɪɚʀɧɚ, 01004 in the yield in the forms of selenomethionine, selenate 2 ȱɧɫɬɢɬɭɬ ɮɿɡɿɨɥɨɝɿʀ ɪɨɫɥɢɧ ɿ ɝɟɧɟɬɢɤɢ ɇȺɇ ɍɤɪɚʀɧɢ (Se(VI)), and in non-protein compounds. On condition ȼɭɥ. ȼɚɫɢɥɶɤɿɜɫɶɤɚ, 31/17, Ʉɢʀɜ, ɍɤɪɚʀɧɚ, 03022 of the application of barium selenate and barium sel- enite during the vegetation period, quinoa seeds accu- Ⱦɥɹ ɡɚɛɟɡɩɟɱɟɧɧɹ ɡɪɨɫɬɚɧɧɹ ɣ ɞɢɜɟɪɫɢɮɿɤɚɰɿʀ ɪɨɫ- mulate organic selenium [67]. ɥɢɧɧɢɰɬɜɚ ɍɤɪɚʀɧɢ ɿ ɪɨɡɲɢɪɟɧɧɹ ɫɭɱɚɫɧɢɯ ɫɿɜɨɡɦɿɧ ɧɚ ɨɫɧɨɜɿ ɭɡɚɝɚɥɶɧɟɧɧɹ ɫɜɿɬɨɜɨɝɨ ɞɨɫɜɿɞɭ ɬɚ ɩɪɨɜɟɞɟɧɢɯ Quinoa is a known photoperiod-responsive crop. ɩɨɥɶɨɜɢɯ ɟɤɫɩɟɪɢɦɟɧɬɿɜ ɞɨɫɥɿɞɠɟɧɨ ɦɨɠɥɢɜɿɫɬɶ ɜɢɪɨ- For instance, cultivars from Ecuador need at least 15 ɳɭɜɚɧɧɹ ɜ ɤɪɚʀɧɿ ɤɿɧɨɚ (Chenopodium quinoa L.). ɇɚɜɟ- days with short illumination period for 10 h to transfer ɞɟɧɨ ɞɚɧɿ ɡ ɫɢɫɬɟɦɚɬɢɱɧɨɝɨ ɩɨɥɨɠɟɧɧɹ ɤɿɧɨɚ ɬɚ ɫɩɨ- to the À owering phase. The duration of the vegetation ɪɿɞɧɟɧɨɫɬɿ ɤɭɥɶɬɭɪɢ ɡ ɿɧɲɢɦɢ ɜɢɞɚɦɢ ɪɨɞɭ Chenopo- period may vary in different years, being from 108 to dium, ɡ ɿɫɬɨɪɿʀ ɞɨɦɟɫɬɢɤɚɰɿʀ ɜɢɞɭ. Ʉɿɧɨɚ ɦɚɽ ɜɢɫɨɤɭ ɯɚɪ- 148 days [68]. ɱɨɜɭ ɰɿɧɧɿɫɬɶ, ʀʀ ɦɨɠɧɚ ɜɢɤɨɪɢɫɬɨɜɭɜɚɬɢ ɭ ɛɟɡɝɥɸɬɟ- ɧɨɜɢɯ ɞɿɽɬɚɯ, ɳɨ ɽ ɜɚɠɥɢɜɢɦ ɤɨɦɩɨɧɟɧɬɨɦ ɪɚɰɿɨɧɭ ɥɸ- Quinoa is uniquely adapted to the cultivation in dif- ɞɢɧɢ. ɉɪɟɞɫɬɚɜɥɟɧɨ ɪɟɡɭɥɶɬɚɬɢ ɜɢɪɨɳɭɜɚɧɧɹ ɤɿɧɨɚ ɭ ferent agroecological regions; it grows at relative hu- 2013í2014 ɪɪ. ɡɚ ɭɦɨɜ ɞɨɫɥɿɞɧɨɝɨ ɫɿɥɶɫɶɤɨɝɨɫɩɨɞɚɪ- midity from 40 to 88 % and can endure the tempera- ɫɶɤɨɝɨ ɜɢɪɨɛɧɢɰɬɜɚ ȱɧɫɬɢɬɭɬɭ ɮɿɡɿɨɥɨɝɿʀ ɪɨɫɥɢɧ ɿ ɝɟɧɟ- tures from í4 °ɋ to +38 °ɋ. This crop uses water with ɬɢɤɢ ɇȺɇ ɍɤɪɚʀɧɢ (ȼɚɫɢɥɶɤɿɜɫɶɤɢɣ ɪɚɣɨɧ Ʉɢʀɜɫɶɤɨʀ ɨɛ- high ef¿ ciency and forms the yield even at 100–200 ɥɚɫɬɿ). Ɍɚɤɢɦ ɱɢɧɨɦ, ɤɭɥɶɬɭɪɚ ɤɿɧɨɚ ɽ ɩɟɪɫɩɟɤɬɢɜɧɨɸ mm of rainfall during the vegetation season [5]. How- ɞɥɹ ɜɿɬɱɢɡɧɹɧɢɯ ɡɟɪɧɨɜɢɪɨɛɧɢɤɿɜ. ȼɜɟɞɟɧɧɹ ɤɿɧɨɚ ɞɨ ever, the physiological mechanisms of stress resistance ɫɿɜɨɡɦɿɧ ɦɨɠɟ ɩɨɤɪɚɳɢɬɢ ɟɤɨɥɨɝɿɱɧɢɣ ɫɬɚɧ ɚɝɪɨɮɿɬɨ- of this crop are studied insuf¿ ciently [69]. ɰɟɧɨɡɿɜ ɬɚ ɫɩɪɢɹɬɢ ɜɿɞɧɨɜɥɟɧɧɸ ɪɨɞɸɱɨɫɬɿ ʉɪɭɧɬɿɜ ɤɪɚʀɧɢ ɛɟɡ ɡɧɢɠɟɧɧɹ ɩɪɢɛɭɬɤɿɜ ɚʉɪɚɪɿʀɜ. In the vegetation season of 2014 with high amount Ʉɥɸɱɨɜɿ ɫɥɨɜɚ: Chenopodium quinoa, ɤɿɧɨɚ, ɛɿɨɥɨɝɿɹ, ɫɢɫ- of rainfall, the ¿ elds of quinoa in Kyiv Region were ɬɟɦɚɬɢɤɚ, ɞɨɦɟɫɬɢɤɚɰɿɹ, ɬɟɯɧɨɥɨɝɿʀ ɜɢɪɨɳɭɜɚɧɧɹ. slightly infested with downy mildew (Peronospora farinosa) and sugarbeet root aphid (Pemphigus fusci- Ʉɢɧɨɚ – ɩɟɪɫɩɟɤɬɢɜɧɚɹ ɞɥɹ ɍɤɪɚɢɧɵ cornis). No infestation with these pathogen and pest «ɡɟɪɧɨɜɚɹ» ɤɭɥɶɬɭɪɚ should be expected under dry conditions of the vegeta- ɋ. Ʌ. Ɇɨɫɹɤɢɧ 1, ȼ. ȼ. ɒɜɚɪɬɚɭ 2 tion season. e-mail: [email protected]; In 2014, the yield of quinoa was 1.05 t/ha, thus mak- e-mail: [email protected] ing the cultivation of this crop highly cost-ef¿ cient. 1 ɂɧɫɬɢɬɭɬ ɛɨɬɚɧɢɤɢ ɢɦ. ɇ. Ƚ. ɏɨɥɨɞɧɨɝɨ ɇȺɇ ɍɤɪɚɢɧɵ According to many authors, the yield may be as high ɍɥ. Ɍɟɪɟɳɟɧɤɨɜɫɤɚɹ, 4, Ʉɢɟɜ, ɍɤɪɚɢɧɚ, 01004 as 2.5 t/ha [70]. 2 ɂɧɫɬɢɬɭɬ ɮɢɡɢɨɥɨɝɢɢ ɪɚɫɬɟɧɢɣ ɢ ɝɟɧɟɬɢɤɢ ɇȺɇ ɍɤɪɚɢɧɵ Therefore, quinoa is a promising crop for domestic ɍɥ. ȼɚɫɢɥɶɤɨɜɫɤɚɹ, 31/17, Ʉɢɟɜ, ɍɤɪɚɢɧɚ, 03022 grain producers. The introduction of quinoa into crop Ⱦɥɹ ɨɛɟɫɩɟɱɟɧɢɹ ɪɨɫɬɚ ɢ ɞɢɜɟɪɫɢɮɢɤɚɰɢɢ ɪɚɫɬɟɧɢɟ- rotation systems can improve ecological conditions ɜɨɞɫɬɜɚ ɍɤɪɚɢɧɵ, ɚ ɬɚɤɠɟ ɪɚɫɲɢɪɟɧɢɹ ɫɨɜɪɟɦɟɧɧɵɯ of agrophytosystems and promote the restoration of ɫɟɜɨɨɛɨɪɨɬɨɜ ɧɚ ɨɫɧɨɜɟ ɨɛɨɛɳɟɧɢɹ ɦɢɪɨɜɨɝɨ ɨɩɵɬɚ soil fertility in the country without diminishing the ɢ ɩɪɨɜɟɞɟɧɧɵɯ ɩɨɥɟɜɵɯ ɷɤɫɩɟɪɢɦɟɧɬɨɜ ɢɫɫɥɟɞɨɜɚɧɚ revenues of farmers and grain producers. The issues ɜɨɡɦɨɠɧɨɫɬɶ ɜɵɪɚɳɢɜɚɧɢɹ ɜ ɫɬɪɚɧɟ ɤɢɧɨɚ (Chenopodi- still to be solved in promoting quinoa cultivation in um quinoa L.). ɉɪɢɜɟɞɟɧɵ ɞɚɧɧɵɟ ɩɨ ɫɢɫɬɟɦɚɬɢɱɟɫɤɨɦɭ Ukraine are the identi¿ cation of crop genotypes opti- ɩɨɥɨɠɟɧɢɸ ɤɢɧɨɚ ɢ ɪɨɞɫɬɜɟ ɤɭɥɶɬɭɪɵ ɫ ɞɪɭɝɢɦɢ ɜɢɞɚ-

8 AGRICULTURAL SCIENCE AND PRACTICE Vol. 2 No. 1 2015 QUINOA AS A PROMISING PSEUDOCEREAL CROP FOR UKRAINE ɦɢ ɪɨɞɚ Chenopodium, ɩɨ ɢɫɬɨɪɢɢ ɞɨɦɟɫɬɢɤɚɰɢɢ ɜɢɞɚ. 12. Takhtajan A. Flowering plants. 2nd ed. Berlin, Springer. Ʉɢɧɨɚ ɢɦɟɟɬ ɜɵɫɨɤɭɸ ɩɢɳɟɜɭɸ ɰɟɧɧɨɫɬɶ, ɟɟ ɦɨɠɧɨ 2009; xlvi + 871 p. ɢɫɩɨɥɶɡɨɜɚɬɶ ɜ ɛɟɡɝɥɸɬɟɧɨɜɵɯ ɞɢɟɬɚɯ, ɱɬɨ ɹɜɥɹɟɬɫɹ 13. Fuentes-Bazan S, Uotila P, Borsch T. A novel phylogeny- ɜɚɠɧɵɦ ɤɨɦɩɨɧɟɧɬɨɦ ɪɚɰɢɨɧɚ ɱɟɥɨɜɟɤɚ. ɉɪɟɞɫɬɚɜɥɟɧɵ based generic classi¿ cation for Chenopodium sensu lato, ɪɟɡɭɥɶɬɚɬɵ ɜɵɪɚɳɢɜɚɧɢɹ ɤɢɧɨɚ ɜ 2013í2014 ɝɝ. ɜ ɭɫɥɨ- and a tribal rearrangement of Chenopodioideae (Cheno- ɜɢɹɯ ɨɩɵɬɧɨɝɨ ɫɟɥɶɫɤɨɯɨɡɹɣɫɬɜɟɧɧɨɝɨ ɩɪɨɢɡɜɨɞɫɬɜɚ ɂɧ- podiaceae). Willdenowia.2012;42:5–24. ɫɬɢɬɭɬɚ ɮɢɡɢɨɥɨɝɢɢ ɪɚɫɬɟɧɢɣ ɢ ɝɟɧɟɬɢɤɢ ɇȺɇ ɍɤɪɚɢ- 14. Aellen P. Chenopodium amaranticolor Coste et Reynier, ɧɵ (ȼɚɫɢɥɶɤɨɜɫɤɢɣ ɪɚɣɨɧ Ʉɢɟɜɫɤɨɣ ɨɛɥɚɫɬɢ). Ɍɚɤɢɦ Ch. purpurascens Jacquin, Ch. giganteum Don, Ch. Qui- ɨɛɪɚɡɨɦ, ɤɭɥɶɬɭɪɚ ɤɢɧɨɚ ɹɜɥɹɟɬɫɹ ɩɟɪɫɩɟɤɬɢɜɧɨɣ ɞɥɹ noa Willd., Ch. Moquinianum Aellen und Ch. Reynieri ɨɬɟɱɟɫɬɜɟɧɧɵɯ ɡɟɪɧɨɩɪɨɢɡɜɨɞɢɬɟɥɟɣ. ȼɜɟɞɟɧɢɟ ɤɢɧɨɚ ɜ Ludwig et Aellen. Eine nomenklatorischen und systema- ɫɟɜɨɨɛɨɪɨɬɵ ɦɨɠɟɬ ɭɥɭɱɲɢɬɶ ɷɤɨɥɨɝɢɱɟɫɤɨɟ ɫɨɫɬɨɹɧɢɟ tischen Studie. Ber. Schweiz. Bot. Ges.1929;38:5–23. ɚɝɪɨɮɢɬɨɰɟɧɨɡɨɜ ɢ ɫɩɨɫɨɛɫɬɜɨɜɚɬɶ ɜɨɫɫɬɚɧɨɜɥɟɧɢɸ ɩɥɨ- 15. Aellen P, Just T. Key and synopsis of the American ɞɨɪɨɞɢɹ ɩɨɱɜ ɫɬɪɚɧɵ ɛɟɡ ɫɧɢɠɟɧɢɹ ɩɪɢɛɵɥɟɣ ɚɝɪɚɪɢɟɜ. species of the genus Chenopodium L. Amer Midl Nat. Ʉɥɸɱɟɜɵɟ ɫɥɨɜɚ: Chenopodium quinoa, ɤɢɧɨɚ, ɛɢɨɥɨɝɢɹ, 1943;30(1):47–76. ɫɢɫɬɟɦɚɬɢɤɚ, ɞɨɦɟɫɬɢɤɚɰɢɹ, ɬɟɯɧɨɥɨɝɢɢ ɜɵɪɚɳɢɜɚɧɢɹ. 16. Aellen P. Chenopodiaceae. G. Hegi. Illustrierte Flora von Mitteleuropa. Berlin & Hamburg: Paul Parey. 1960– REFERENCES 1961; Band 3, Teil 2 (Lief. 2–4):533–762. 1. Rosegrant M, Ringler C, Sinha A, Huang J, Ahammad H, 17. Scott AJ. A review of the classi¿ cation of Chenopodium Zhu T, Msangi S, Sulser T, Batka M. Exploring alterna- L. and related genera (Chenopodiaceae). Bot Jahrb Syst. tive futures for agricultural knowledge, science and tech- 1978;100:205–20. nology (AKST). ACIAR Project Report ADP/2004/045. 18. Mosyakin SL An outline of a system for Chenopodium L. Canberra, ACIAR.2009;84 p. (species of Europe, North and Central Asia). Ukr Bot Zh. 2. Food Outlook: Biannual report on global food markets. 1993; 50(5):71–7. June 2013. Rome, FAO.2013;132 p. 19. Mosyakin SL, Chenopodium L Flora Europae Orientalis. 3. Neglected crops: 1492 from a different perspective. Eds J. St. Petersburg, Mir i Semya-95.1996;Vol. 9:27–44. E. Hernandes Bermejo, J. Leon. FAO Plant Production and 20. Mosyakin SL The system and phytogeography of Che- Protection Series, no. 26. Rome, FAO.1994; xxii + 341 p. nopodium subgen. Blitum (L.) I. Hiitonen (Chenopodia- 4. Descriptores para quinua (Chenopodium quinoa Willd.) ceae). Ukr Bot Zh.2002;59(6):696–701. y sus parientes silvestres. Roma, Bioversity Internation- 21. Mosyakin SL The system and phytogeography of Cheno- al. 2013; vi + 52 p. podium subgen. Chenopodium (Chenopodiaceae). Ukr 5. Quinoa: An ancient crop to contribute to world food se- Bot Zh.2003;60(1):26–32. curity. FAO, Regional Of¿ ce for Latin America and the 22. Mosyakin SL, Clemants SE. New nomenclatural combi- Caribbean. 2011; vi + 55 p. nations in Dysphania R. Br. (Chenopodiaceae): taxa oc- 6. Bazile D, Fuentes F, Mujica A. Historical perspectives curring in North America. Ukr Bot Zh.2002;59(4):380– and domestication. Ch. 2. Quinoa: botany, production 85. and uses. Eds A. Bhargava, S. Srivastava. Wallingford, 23. Mosyakin SL, Clemants SE. Further transfers of glan- CABI.2013;16–35. dular-pubescent species from Chenopodium subg. Am- 7. Schlick G, Bubenheim DL. Quinoa: candidate crop for brosia to Dysphania (Chenopodiaceae). J Bot Res Inst NASA’s Controlled Ecological Life Support Systems. Texas.2008;2(1):425–31. Progress in new crops. Ed. J. Janick. Arlington, ASHS 24. Clemants SE, Mosyakin SL. Dysphania R. Brown; Che- Press.1996;632–40. nopodium Linnaeus. Flora of North America north of 8. The Angiosperm Phylogeny Group. An update of the An- Mexico. Ed. by FNA Editorial Committee. New York, giosperm Phylogeny Group classi¿ cation for the orders Oxford Univ. Press.2003;Vol. 4:267–99. and families of À owering plants: APG III. Bot J Linnean 25. Zhu Gelin, Mosyakin SL, Clemants SE. Chenopodiaceae. Soc. 2009; 161(2):105–21. Flora of China. Eds Wu Zhengyi, P. H. Raven. Beijing, 9. Mosyakin SL. Families and orders of angiosperms of the Science Press & St. Louis, Missouri Botanical Garden À ora of Ukraine: a pragmatic classi¿ cation and placement in Press.2003;Vol. 5:351–414. the phylogenetic system. Ukr Bot Zh.2013;70(3):289–307. 26. Kadereit G, Borsch T, Weising K, Freitag H. Phylogeny of 10. The Angiosperm Phylogeny Group. An ordinal classi¿ ca- Amaranthaceae and Chenopodiaceae and the evolution of

tion for the families of À owering plants. Ann Missouri C4-photosynthesis. Int J Plant Sci.2003;164(6):959–986. Bot Gard.1998;85(4):531–53. 27. Kadereit G, Gotzek D, Jacobs S, Freitag H. Origin and 11. The Angiosperm Phylogeny Group II. An update of the age of Australian Chenopodiaceae. Org Divers Evol. Angiosperm Phylogeny Group classi¿ cation for the or- 2005;5(1):59–80. ders and families of À owering plants: APG II. Bot J Lin- 28. Fuentes-Bazan S, Mansion G, Borsch T. Towards a nean Soc.2003;131(4):399–436. species level tree of the globally diverse genus Che-

AGRICULTURAL SCIENCE AND PRACTICE Vol. 2 No. 1 2015 9 MOSYAKIN et al. nopodium (Chenopodiaceae). Mol Phylogenet Evol. 46. Baschilov VA. Appearance of crop plants in the old- 2012;62(1):359í74. est agricultural centers of America. Latin America. 29. Mosyakin SL. New nomenclatural combinations in Bli- 1980;(5):92–101. tum, Oxybasis, Chenopodiastrum, and Lipandra (Cheno- 47. Bashilov VA. Periodisation and tempos of the historical podiaceae). Phytoneuron.2013;2013-56:1–8. process of the in the Middle East 30. Mosyakin SL, Clemants SE. New infrageneric taxa and and in the New World. Ancient civilizations of the Orient. combination in Chenopodium L. (Chenopodiaceae). No- Tashkent, FAN Publ.1986;61–6. von. 996; 6(4):398–403. 48. Schnirelman VA. Ideas of N. I. Vavilov and modern 31. Mosyakin SL. Teloxys Moq., Dysphania R. Br., Blitum L., data on formation of early centers of food production. Chenopodium L. Conspectus Florae Europae Orientalis. Vavilov’s heritage in modern biology. Moscow, Nauka. Eds N. N. Tzvelev, D. V. Geltman. St Petersburg, Mos- 1989;299–317. cow, KMK Scienti¿ c Press.2012;Vol. 1:276–86. 49. Prehistoric food production in North America. Ed. R. I. 32. Sukhorukov AP, Zhang M. Fruit and seed anatomy of Ford. Ann Arbor, Univ. Michigan Press.1985;411 p. Chenopodium and related genera (Chenopodioideae, 50. Diamond J. Guns, germs and steel. A short history of Chenopodiaceae/Amaranthaceae): Implications for evo- everybody for the last 13,000 years. London, Vintage. lution and taxonomy. PLoS One.2013;8(4):e61906. 1998;480 p. 33. Crawford DJ, Wilson HD. Allozyme variation in sev- 51. Diamond J. Evolution, consequences and future of eral closely related diploid species of Chenopodium in plant and animal domestication. Nature.2002;418(6898): the Western United States. Amer J Bot. 1979;66(3):237– 700–7. 44. 52. Smith BD. Chenopodium as a prehistoric domesticate 34. Wilson HD. Arti¿ cial hybridization among species of in eastern North America: Evidence from Russell Cave, Chenopodium sect. Chenopodium. Syst Bot.1980;5(3): Alabama. Science.1984;226(4671):165–7. 253–63. 53. Smith BD. Eastern North America as an independent cen- 35. Wilson HD. Genetic variation among South American ter of plant domestication. Proc Natl Acad Sci U S A. populations of tetraploid Chenopodium sect. Chenopo- 2006;103(33):12223–8. dium subsect. Cellulata. Syst Bot.1981;6(4):380–98. 54. Hunziker AT. Estudios sobre Amaranthus. I. Los espe- 36. Wilson HD. Domesticated Chenopodium of the Ozark cies alimenticias de Amaranthus y Chenopodium culti- Bluff dwellers. Econ Bot.1981;35(2):233–9. vadas por los indios de América. Revista Argent. Agron. 37. Wilson HD. Chenopodium quinoa Willd.: variation and 1943;10:297–354. relationships in southern South America. Natl Geogr Soc 55. Hunziker AT. Los pseudocereales de la agricultura in- Res Reports.1985;19:711–21. dígena de América. (Mus. Bot. Córdoba; Publ. Misc. 38. Wilson HD. Allozyme variation and morphological re- No. 3.) Buenos Aires, ACME Agency.1952;104 p. lationships of Chenopodium hircinum (s.l.). Syst Bot. 56. Sauer JD. The grain amaranths: a survey of their 1988;13(2):215–28. history and classi¿ cation. Ann Missouri Bot Gard. 39. Wilson HD. Quinoa biosystematics I: Domesticated pop- 1950;37(4):561–632. ulations. Econ Bot. 1988;42(4):461–77. 57. Sauer JD. The grain amaranths and their relatives: a re- 40. Wilson HD. Quinoa biosystematics II: Free-living popu- vised taxonomic and geographic survey. Ann Missouri lations. Econ Bot. 1988;42(4):478–94. Bot Gard.1967; 54(2):103–37. 41. Wilson HD. Quinoa and relatives (Chenopodium sect. 58. Sauer CO. Agricultural origin and dispersal. Cambridge, Chenopodium subsect. Cellulata). Econ Bot.1990; MTI Press.1969;143 p. 44(3):92–110. 59. Mosyakin SL, Robertson KR. Amaranthus Linnaeus (Am- 42. Wilson HD, Heiser CB. The origin and evolutionary re- aranthaceae). Flora of North America north of Mexico. lationships of “huazontle޵ (Chenopodium nuttaliae Saf- Ed. by FNA Editorial Committee. New York, Oxford ford), domesticated chenopod of Mexico. Amer J Bot. Univ. Press.2003;Vol. 4:410í35. 1979;66(2):198–206. 60. Ford-Lloyd BV, Williams JT. A revision of Beta section 43. Lack HW, Fuentes S. The discovery, naming and typi¿ ca- Vulgares (Chenopodiaceae), with new light on the origin tion of Chenopodium quinoa (Chenopodiaceae). Willde- of cultivated beets. Bot J Linnean Soc.1975;71(2):89–102. nowia.2013;43(1):143–9. 61. de Bock TSM. The genus Beta: domestication, taxonomy 44. Mosyakin SL. Life strategies of wild relatives of crop and interspeci¿ c hybridization for plant breeding. Acta plants as prerequisites of their domestication. Botany and Hort. 1986;182:335–43. mycology: Modern horizons. In memory of Academician 62. van Dijk H, Desplanque B. European Beta: Crops and A. M. Grodzinsky (1926–1988. Kyiv, Akademperiodyka. their wild and weedy relatives. Plant evolution in man- 2007;150–68. made habitats. Proc. VII Int. IOPB Symp. (Amsterdam, 45. Vavilov NI. Centers of origin of crops. Leningrad, All-Union The Netherlands; August 10í15, 1998). Amsterdam, Inst. of Applied Botany and New Crops.1926;248 p. Univ. Amsterdam Press.1999;257–70.

10 AGRICULTURAL SCIENCE AND PRACTICE Vol. 2 No. 1 2015 QUINOA AS A PROMISING PSEUDOCEREAL CROP FOR UKRAINE 63. Moerman DE. Native American ethnobotany. Portland, Sciences Georg-August-University of Gottingen. Gottin- Oregon, Timber Press. 1998;927 p. gen. 2004;79 p. 64. Jellen EN, Kolano BA, Sederberg MC, Bonifacio A, 67. Kitaguchi T, Ogra Y, Iwashita Y, Suzuki KT. Speciation Maughan PJ. Chenopodium. Wild crop relatives: genom- of selenium in selenium-enriched seeds, buckwheat ic and breeding resources. Ed. C. Kole. Berlin, Springer. (Fagopyrum esculentum Moench) and quinoa (Che- 2011;35–61. nopodium quinoa Willdenow). Eur Food Res Technol. 2008;227(5):1455–60. 65. Jellen EN, Maughan PJ, Fuentes F, Kolano BA. Capi- 68. Jacobsen SE. The worldwide potential for quinoa (Cheno- tulo 1.1: Botánica, ¿ logenia y evolución. Estado del arte podium quinoa Willd.). Food Rev Int. 2003;19(1í2):167– de la quinua en el mundo en 2013. Eds D. Bazile et al. 77. Santiago de Chile, FAO y CIRAD (Montpellier, Francia). 69. Tapia ME. Cultivos andinos subexplotados y su aporte a 2014;12–25. la alimentación [Underexploited Andean crops and their 66. Thanapornpoonpong S. Effect of nitrogen fertilizer on nutritional contribution]. 2nd ed. Santiago (Chile), FAO. nitrogen assimilation and seed quality of amaranth (Ama- 1997;273 p. ranthus spp.) and quinoa (Chenopodium quinoa Willd.). 70. Jacobsen SE. Adaptation of quinoa (Chenopodium qui- Doctoral Dissertation submitted for the degree of Doctor noa) to Northern European agriculture: studies on devel- of Agricultural Sciences of the Faculty of Agricultural opmental pattern. Euphytica.1997;96(1):41–8.

AGRICULTURAL SCIENCE AND PRACTICE Vol. 2 No. 1 2015 11