Perennial Grains in Russia: History, Status, and Perspectives

Abstract

The review summarizes the historical and current research on perennial grain breeding in Russia within the context of growing global interest in perennial crops. N.V. Tsitsin’s pioneering work in the 1930s produced the first wheat–wheatgrass amphiploids, which demonstrated the capacity to regrow after harvest and survive for 2–3 years. Subsequent research at the Main Botanical Garden in Moscow focused on characterizing Tsitsin’s material, selecting superior germplasm, and expanding genetic diversity through new cycles of hybridization and selection. This work led to the development of a new crop species, Trititrigia, and the release of cultivar ‘Pamyati Lyubimovoy’ in 2020, designed for dual-purpose production of high-quality grain and green biomass. Intermediate wheatgrass (Thinopyrum intermedium) is native to Russia, where several forage cultivars have been released and cultivated. Two large-grain cultivars (Sova and Filin) were developed from populations provided by the Land Institute and are now grown by farmers. Perennial rye was developed through interspecific crosses between Secale cereale and S. montanum, demonstrating persistence for 2–3 years with high biomass production and grain yields of 1.5–2.0 t/ha. Hybridization between Sorghum bicolor and S. halepense resulted in two released cultivars of perennial sorghum used primarily for forage production under arid conditions. Russia’s agroclimatic diversity in agricultural production systems provides significant opportunities for perennial crop development. The broader scientific and practical implications of perennial crops in Russia extend to climate-resilient, sustainable agriculture and international cooperation in this emerging field.

1. Global Tendency for Breeding and Production of Perennial Grains

The current scientific concept of perennial grains has been primarily associated with the Land Institute (Kansas, USA) and its founder Wes Jackson [1], who presented a vision of perennial agriculture in his book “New Roots for Agriculture”. The book argued that industrialization of agriculture has had severe effects on nature and society. A root cause is the dominance of annual plants cultivated in monocultures. Annual crops require yearly clearing of vegetation, resulting in soil erosion and other forms of ecosystem degradation. Monocultures are susceptible to agricultural pests and weeds. By contrast, perennial polycultures informed by natural ecosystems promise more sustainable agroecosystems with the potential to revitalize the economic foundation of farming and rural societies [2].

The global context of perennial grain development has been reviewed by Olsson et al. [3], who listed four main opportunities for the new production system: (a) perennial polycultures appeal to farmers; (b) challenge to contemporary agricultural and food industries; (c) growing priority of soil health; (d) interest in public funding. The authors believe that the time is ripe for the onset of a perennial revolution. The goal of this revolution is the rapid development of entirely new, high-yielding perennial grain crops that can replace the current repertoire of annuals. The result will be cropping systems that preserve soil, store carbon efficiently, require minimal inputs in terms of commercial energy and machinery, utilize available water effectively, are increasingly self-sufficient in nitrogen, and can unlock stores of phosphorus in agricultural soils.

Perennial grains represent staple cereals (wheat, barley, rye, rice, maize, sorghum) with the capacity to grow for several seasons in the same field after one planting [4]. The main advantages of perennial grains include their reduced negative environmental impact, contribution to soil health, mitigation of climate change, and support of biodiversity. Economically viable perennial grain production is based on reduced costs associated with annual soil preparation and planting, reasonable yield and suitable grain quality, and ecological services. A fundamental difference between annual and perennial plants is the root system [3]. Annuals develop roots for just one season, whereas perennials build and accumulate typically deeper and wider root systems over numerous years. From a farming perspective, perennial grains could be described as a farmer’s dream—a cultivar that is planted once and then harvested every season for several years with minimal land management in between. Instead of 4–10 tractor passes per year as with annual cultivars, only 1–5 passes for harvesting and nutrient, pest, and weed management would be required. From a farm economics perspective, farming perennial grains could translate into a significant reduction in production costs, resulting in increased total factor productivity.

Comparative physiology, biology, ecology, and genetics of perennial and annual cereals was reviewed by Chapman et al. [5]. Perennialism is a highly complex trait, unlikely to be conferred by a single gene. Ancestral state reconstructions using phylogenetic approaches demonstrated that annual species are generally derived from perennial ancestors, often via multiple independent events within a genus. Two main directions dominate in the development of modern perennial grains: (a) domestication of perennial grasses with potential to become economically viable grain crops (Kernza–Thinopyrum intermedium syn. Agropyron glaucum syn., and Elytrigia intermedia as alternatives to perennial wheat, and Hordeum bulbosum to perennial barley); (b) crosses of annual crops with perennial wild relatives to combine productivity and perennity (wheat, rye, sorghum, maize, and rice). Both directions have been pursued globally, and some examples are given below.

Perennial wheat has been proposed as an alternative crop in the early 1930s by the Russian scientist Nikolay Tsitsin, who started systematic crosses between wheat and perennial grasses to develop new crops [6]. His efforts resulted in stable wheat amphiploids with some degree of perennity as well as the successful transfer of genes from wild wheat relatives to cultivated varieties. Work on perennial wheat breeding was also conducted in the USA, Australia, China, and Europe using similar approaches of combining common wheat genomes with various perennial grasses. However, this work did not develop into a commercial product.

As global societal environmental concerns grew over time, the development of staple perennial food crops and production systems gained popularity with several research and breeding programs developing commercial products. Intermediate wheatgrass (IWG), or Kernza (Thinopyrum intermedium), went through robust domestication processes for grain size and productivity and now represents a commercial crop in North America [7]. Since initial work in the 1980s, breeding programs have been initiated in Kansas, Minnesota, and Utah (United States), Manitoba (Canada), and Uppsala (Sweden). Coupled with advances in molecular technologies, many of these programs have harnessed the power of genomic selection and other cutting-edge tools to rapidly improve IWG. This has resulted in estimated gains of up to 8% per year for spike yield, and across eight breeding cycles, grain yield has increased 9% per cycle. Nutritional quality studies demonstrated that IWG had 50% higher protein, 129% higher dietary fiber, and 65% higher ash content than the reference whole wheat flour [8]. Calcium and selenium were 267% and 492% higher, respectively, in IWG than whole wheat flour. The results support potential benefits of Kernza for human nutrition. Enrichment of wheat flour with IWG grain significantly affected gluten content and quality as well as bread characteristics [9]. IWG flour addition had a positive effect on phenolic and antioxidant properties. These results indicated that IWG has great potential to be used in bread production as a novel, healthy, and sustainable ingredient.

Perennial rice was developed through crossing annual Asian rice (Oryza sativa) with its perennial African relative Oryza longistaminata [10]. From a single planting, irrigated perennial rice produced grain for eight consecutive harvests over four years, averaging 6.8 t/ha per harvest versus 6.7 t/ha of replanted annual rice, which required additional labor and seeds. Four years of cropping with perennial rice resulted in soils accumulating 0.95 t/ha of organic carbon and 0.11 t/ha of nitrogen per year, along with increases in plant-available water capacity (7.2 mm). Perennial cultivars are strongly preferred by farmers; growing them saves 58% of labor and 49% of input costs in each regrowth cycle. Suited to a broad range of frost-free environments between 40° N and 40° S, perennial rice is a step change with potential to improve livelihoods, enhance soil quality, and inspire research on other perennial grains.

Domestication efforts are underway to develop perennial food oil crops including sunflower (Helianthus tuberosus, H. maximiliani), silflower (Silphium integrifolium) [11], and flax [12]. Sainfoins (Onobrychis spp.), perennial legumes domesticated as ancient forages, have been bred as food crops with Baki™ bean tradename [13]. Perennial sorghum demonstrated high potential for sustainable cropping systems in Africa [14].

The importance of perennial crops was exemplified by an International Workshop “Pathways to a Perennial Future”, which took place in Mexico in March 2025. The workshop was attended by over 100 scientists from 25 countries. The scientific agenda was focused on perennial crop improvement and genetics, sustainable production practices, consumption, and marketing. The workshop contributed to the development of research directions and global cooperation framework. The workshop working group on perennial wheat summarized the status of breeding and research globally and established an informal network. The group realized that little knowledge is available on post-Tsitsin perennial wheat breeding and research in Russia. This includes the recent advances and performance of the wide crosses program at the Main Botanical Garden named after N.V. Tsitsin in Moscow as well as evaluation and use of global perennial wheat collection in Western Siberia (city of Omsk) earlier reported by Hayes et al. [15]. Thus, the objective of the current review is to close this knowledge gap by presenting the results and perspectives of perennial wheat breeding and research in Russia, with the main focus being on the last 20–30 years. As the review was being written, relevant information on other perennial grains became available and the focus of this document expanded to include the genetic resources of rye, IWG, and sorghum.

2. Russia Crop Production and Potential for Perennial Grains

Comprehensive coverage of Russian agricultural production and trade is presented in a relatively recent book titled “Russia’s Role in the Contemporary International Agri-Food Trade System” [16]. The book preface lists two major developments in Russian agriculture: remarkable improvement in agricultural production that Russia has experienced since 2004 and commitment to extend food policy beyond food security and self-sufficiency to become a food exporter. The emergence of Russia as a major grain exporter is one of the most remarkable storylines in the past decade.

The total arable land of Russia exceeded 80 million hectares in 2024. The entire country is divided into nine federal regions with specific agroclimatic conditions, soils, and respective differences in cropping systems (

Table 1. Crop area across Russia federal regions in 2024 (source: http://ssl.rosstat.gov.ru/storage/mediabank/posev-4cx_2024.xlsx (accessed on 15 May 2025)).

Crop	Crop Area in Federal Regions, M ha
	Central	North-West	South	North Caucasus	Volga	Ural	Siberia	Far East	Total
Arable land	15.7	1.3	13.3	4.4	24.1	5.0	14.2	2.1	80.2
Winter cereals	3.2	0.1	6.3	2.3	4.6	0	0	0	17.5
Spring wheat	1.3	0	0.2	0	3.6	2.2	4.9	0.2	12.4
Spring barley	1.3	0.1	0.5	0.1	2.4	0.7	1.2	0.1	6.3
Maize for grain	0.9	0	0.7	0.5	0.4	0	0	0.2	2.6
Pulses	0.4	0	0.6	0.3	1.4	0.2	0.9	0	3.9
Oil crops	4.3	0.1	3.1	0.5	5.8	0.6	3.1	1.2	18.9
Potato	0.2	0	0.1	0.1	0.2	0.1	0.1	0.1	1.0
Forage crops	2.8	0.8	0.6	0.3	4.4	0.9	2.3	0.2	12.3

). Winter and spring wheat collectively occupied 28.5 million hectares or 35.5% of the total planted area in 2024. In fact, the share of winter and spring wheat reflects the agroclimatic conditions in specific regions. The South and North Caucasus regions almost entirely grow winter wheat and represent the country’s breadbasket. Eastern regions (Ural, Siberia, and the Far East) are dedicated to spring wheat as winter is too cold and severe for sustainable winter wheat production. In the two largest regions (Central and Volga), spring wheat is gradually being replaced by winter wheat due to favorable effects of climate change (Abys et al. 2024) [17]. The second most widely grown crop is sunflower (9.8 million hectares) followed by barley (6.9 million hectares) and soybeans (4.3 million hectares). Dry peas dominate among pulses with 2.2 million hectares. Rapeseed (2.7 million hectares) and flax (1.7 million hectares) complement sunflowers in the oil crops category. The yield of most crops is below potential and behind the yields of similar ecologies in North America: wheat yielded 2.89 t/ha and sunflower 1.72 t/ha in the favorable 2024 season.

So far, perennial grains have not been cultivated in Russia on a commercial scale, with the exception of IWG cultivar Sova with an estimated area of 5000 ha. Even this cultivar is mainly grown for forage, and the market for its grain has not been developed yet. However, Russia’s diversity of agroclimatic zones and cropping systems represents an opportunity for perennial cereals. The South and North Caucasus regions may be well suited for perennial wheat, combining mild winters and sufficient rainfall year-round. In the Central region, yearly precipitation is high, but winter may be too cold for continuous perennial wheat survival. In the Volga region, in addition to cold winters, summer drought may affect perennial crops during the critical stage between harvest and the next overwintering. Total rice area in Russia was 0.2 million hectares in 2024, primarily planted in the North Caucasus (Krasnodar region). However, winter temperatures may fall to −15 to −20 °C, making perennial rice risky for cultivation unless cold tolerance is incorporated. Rye bread is traditionally consumed in Russia, with the crop area exceeding 0.65 million hectares in 2024. Provided that perennial rye would match winter rye in grain quality, it can be a viable alternative. The area under sorghum for grain is only 37,000 ha in the Southern and Volga regions, and its perennial cultivars can be tested and evaluated for cultivation potential across the country. Sainfoin is a popular forage crop and can be used for food once the concept is confirmed, relevant commercial cultivars are available, and marketing justifies production. Overall, the farming community is open to innovations and is aggressively searching for new cultivars and technology. The lack of research and knowledge on perennial grains limits their conceptual consideration and possible acceptance in the country.

3. Legacy of Nikolay Tsitsin (1898–1980)

A recent publication by Goncharov [6] provides an overview of Nikolay Tsitsin’s biography and scientific achievements. He was born near Saratov in the Volga region in 1898. He graduated from Saratov State Agricultural Institute in 1927 and defended his thesis on segregation of crosses between bread and durum wheat. The idea of crossing wheat and Agropyron came to him when he saw a winter wheat field damaged by harsh conditions, with only a few bright and green spots being wheatgrass. The first crosses between bread wheat and glaucous wheatgrass (Agropyron glaucum) were obtained in 1930 in the Rostov region where he worked at a large-grain farm. In this review, we commonly use the species name Thinopyrum intermedium, which is the same species as Agropyron glaucum and Elytrigia intermedia (both more commonly used in Russia).

In 1932, N.V. Tsitsin moved to Omsk in Western Siberia as the head of the laboratory of wheat–wheatgrass hybrids of the Siberian Research Institute of Grain Farming, eventually being promoted to the director position. In Omsk, N.V. Tsitsin [18] significantly expanded the research, successfully crossing a number of species of wheat and wheatgrass and starting their comprehensive selection study. The main target was the development of winter-hardy winter wheat suitable for Siberian conditions. N.V. Tsitsin began to widely use another property of wheatgrass in breeding–perennity. He was the first in the world to create a new cereal crop–perennial wheat [19]. He named it Triticum agropyrotriticum Cicin [20], ultimately distinguishing two subspecies: T. agropyrotriticum ssp. perenne Cicin (perennial wheat) and T. agropyrotriticum ssp. submittans Cicin (regrowing wheat) [21]. Constant forms of perennial wheat were obtained that yielded for three years after sowing. Later, Tsvelev [22] described this genus as Trititrigia cziczinii Tzvelev. Currently, the crop is included in the State Register of Breeding Achievements [23].

In 1940–1949, N.V. Tsitsin was the director of the Institute of Grain Farming of the Central Regions of the Non-Black Soil Zone in Moscow and the head of the laboratory, in which work on remote hybridization was widely carried out [6]. In 1938–1949 and 1954–1957, N.V. Tsitsin was the director of the All-Union Agricultural Exhibition in Moscow. At the same time, he was responsible for the establishment of the Main Botanical Garden (MBG) of the Academy of Sciences in Moscow, which opened to the public in 1959. The Department of Wide Crosses of MBG became the center of hybridization and breeding work, conducting field experimentation at a station in Snegiri, 40 km west of Moscow. The department continues its work until the present time.

Numerous successful syntheses of new plant forms allowed Tsitsin to formulate a theory of speciation in the plant kingdom, according to which, new constant species appear through a series of temporary unstable forms [21]. Despite the fact that representatives of the tribe Triticeae are a convenient object for the study of interspecific hybridization, the accompanying act of polyploidization, as well as overcoming sterility in interspecific and intergeneric F₁ hybrids, required the solution of issues related to incompatibility. Another significant problem was the lack of a favorable combination of traits in newly created hybrid forms. This required the development of specific methods that would consistently have a positive effect. N.V. Tsitsin spent many years solving these difficult issues.

N.V. Tsitsin’s pioneering work on wheat wide crosses involved two main species of wheatgrass: Elytrigia intermedia (Host) Nevski (syn. Thinopyrum intermedium) and Elytrigia elongata (Host) Nevski (syn. Agropyron elongatum (Host) P.Beauv.) [24]. Amphiploids of wheat and wheatgrass (2n = 56) belonged to two groups: perennial and regrowing. The work on perennial wheat never resulted in commercial cultivars. Breeding of regrowing wheat was prioritized and produced several cultivars officially registered in Russia including Otrastayushaya (Regrowing) 38 and Pamyati Lyubimovoy. Both are stable 56-chromosome Trititrigia amphiploids. The general scheme of crosses and selection to develop Trititrigia and wheat–wheatgrass hybrids (WWHs) is presented in Figure 1.

The main practical outcome came from wheat–wheatgrass hybrids, which were developed through top-crosses or backcrosses to wheat, represented wheat types and possessed wild species genetic material. Six spring wheats were released and widely cultivated in newly developed virgin lands of Western Siberia and Northern Kazakhstan in the 1960s. Cultivar Botanicheskaya 2, obtained from crossing Mexican variety Pitic 62 with WWH Raduga, was widely cultivated in the Urals, Siberia, and Northern Kazakhstan. It was resistant to leaf rust and provided yields of 3 t/ha and more [25]. Cultivar Grekum 114 [21] was resistant to loose smut, powdery mildew, shattering, and lodging; was characterized by high content of protein and gluten; and had excellent baking properties. In addition to spring wheat, several winter wheat cultivars (PPG 599, PPG 1, PPG 186) were released in the 1950s–1960s and cultivated in the Central region of Russia. Perennial line M2 served as the main parent for developing winter wheat cultivars. They were characterized by high winter hardiness and allowed the expansion of winter wheat cultivation area to regions normally sown with rye.

4. Trititrigia as a Man-Made Crop from Wheat–Wheatgrass Crosses

The Department of Remote Hybridization was part of the Moscow Botanical Garden of the Academy of Sciences. After the decision to build the Main Botanical Garden, it became part of the new Garden. In connection with the large-scale expansion of breeding work organized by N.V. Tsitsin, experimental farm “Snegiri” (Moscow region) was established in 1953. It was here that large-scale work was launched on a broad study of the genetic basis, biology, and morphology of germplasm developed by N.V. Tsitsin and his colleagues. He continued to directly supervise the research work on the creation of perennial wheat. After the death of N.V. Tsitsin in 1980, his colleague V.F. Lyubimova continued his work. However, even during the lifetime of Nikolay Vasilyevich, due to the labor intensity of work on the study of perennity, the vector of research was shifted to the study of the ability of perennial wheat to regrow. It is the regrowing forms that are the basis of the genetic resources collection of Trititrigia lines, which are preserved in the department to this day.

A summary of the state of the collection of intermediate wheat–wheatgrass hybrids 30 years after Tsitsin was presented by Upelniek et al. [19]. The collection of Trititrigia was represented by more than 250 forms developed through multiple complex crosses between the original WWHs, involving many released varieties of bread and durum wheat. Modern hybrids have great variation in the length of the spike and its morphology (Figure 2). The longest loose spikes were 20–27 cm long, with 20–22 spikelets and 4–5 flowers per spikelet. One of the important achievements in the last 30 years was the development of stable forms according to the main morphological and biological characteristics. All the modern germplasm is strictly self-pollinated. Many of them have been grown in the same field for more than 20 years with regular replanting while retaining phenotypic identity.

The chromosomal composition of four WWH cultivars (Zernokorovaya 169, Istra-1, Ostankinskaya, and Otrastayashchaya 38) was studied by the method of genomic in situ hybridization [26]. Differentiation of wheatgrass chromosomes was revealed as well as differentiation, centromeric index, and absolute size differences between the cultivars. The presence of telosomes and translocations indicated an ongoing evolution process, making these cultivars a promising resource for obtaining introgressions of wheatgrass genes. Cultivar Ostankinskaya, representing 56-chromosome Trititrigia, was officially released in 1991 (https://graingenes.org//GG3/node/276 accessed 15 May 2025). It was included in a network of 21 experiments across nine countries on four continents to evaluate the relative performance of early generation perennial cereal material [15]. The overall performance index included grain size in year 1, total grain yield over the experimental period (1–3 years), and persistence (frequency in years 2 and 3), with Ostankinskaya being ranked #1 across sites with cold winters.

Modern representatives of the WWH have been developed through complex intervarietal and interhybrid crosses between a large number of modern varieties of winter bread and durum wheat and three species of wheatgrass that have undergone multiple selections [19]. These genetic resources can be divided into three groups: (1) perennial (No. 4015, 1760, 548, M3202, etc.), which have stable, good regrowth in the year following harvesting with 30–60 plants per square meter; (2) regrowing material, which does not survive the next year after grain harvesting (No. M12, 4082, ZP26)—i.e., they are almost annual but with good regrowth of green biomass after grain harvesting, which can be mowed 3–4 times during the growing season; 3) intermediate (No. M209, 2087, 4044, 5542, and many others) with variable numbers of overwintered plants in the second year depending on winter conditions and duration of vernalization stage. This latter group is the most numerous.

Genetic diversity of 24 representatives of two T. cziczinii subspecies (ssp. submitans and ssp. perenne) and 17 genotypes of Triticum aestivum, Triticum durum, Agropyron glaucum, and Agropyron elongatum was studied using 224 AFLPs (Amplified Fragment Length Polymorphism) [27]. The results indicated a greater genetic relatedness of T. cziczinii to T. aestivum than to representatives of the genus Agropyron. According to the cluster analysis, representatives of T. cziczinii and varieties of bread wheat were combined into a single subcluster, within which the samples of two species formed separate groups. At the same time, the evaluation of the intraspecific genetic diversity of T. cziczinii showed no reliable differentiation of representatives of the subspecies submitans (regrowing) and perenne (perennial), which is probably due to the uncertain genetic nature of perennity.

WWH germplasm was extensively studied for agronomic traits. Kocheshkova et al. [28] evaluated variability of 87 wheat–wheatgrass hybrids (bred by N.V. Tsitsin, V.F. Lyubimova, and V.I. Belov) for resistance to pre-harvest sprouting. Most entries showed susceptibility, which may pose a serious problem. Although all evaluated entries had red grain associated with higher seed dormancy, only a few showed relatively high resistance. Using PCR-based markers, polymorphism of the haplotypes of the wheatgrass ThVp-1 gene was identified and revealed significant effects on pre-harvest sprouting resistance. Salinity tolerance was assessed in 10 WWHs based on seedling reaction to variable NaCl concentrations [29]. Three genotypes (2087, 4082, 548) demonstrated relatively high degrees of tolerance. Adult plant leaf rust resistance to 10 pathotypes collected across Russia was evaluated in 18 octoploid WWHs [30]. All WWHs possessed chromosomes from Agropyron S, J, and Js sub-genomes. Seven genotypes including cultivar Istra showed high degree of resistance to all pathotypes and represent valuable genetic resources for breeding.

Long-term data is available on grain and biomass productivity of Trititrigia. Ivanova et al. [31] summarized field trial results from 2013 to 2018, comparing six Trititrigia lines with winter wheat (Rubezhnaya) and rye (Snegirevskaya). On average, dry hay yield for WWH varied from 12.3 (line ZP 548) to 13.7 t/ha (line ZP 1692), while the yield of winter wheat and rye hay was below 7.7 t/ha. The grain yield of Trititrigia ranged from 2.7 (ZP 548) to 3.2 t/ha (M 3202), with winter wheat yielding 4.0 t/ha and Snegirevskaya winter rye yielding 4.4 t/ha. Trititrigia lines were recommended for the following: (1) cultivation of grain and their subsequent mowing or free grazing; (2) growing of green fodder with obtaining two or three harvests per field season. Grain productivity of WWH genotypes was associated with different yield components including spike productivity and 1000-kernel weight [24]. Materials with longer spikes tended to produce lower yields.

In another study, Kvitko et al. [32] evaluated dynamics of biomass growth of Trititrigia compared to winter rye Snegirevskaya. At the booting stage, rye accumulated 23.5 t/ha of biomass compared to 22.3 t/ha for Trititrigia ZP 26. At the stage of milk maturity, the biomass yield was, respectively, 37.4 and 35.4 t/ha. However, winter rye did not recover biomass after harvest, while Trititrigia produced an additional 18.8 t/ha.

The characteristic feature of Trititrigia grain is its exceptional quality, surpassing the grain of bread wheat [33]. The grain of Trititrigia had the following characteristics: crude protein—18.2–19.1%, crude fat—1.3%, crude fiber—2.8–5.0%. The exchange energy of grain is 11.7–12.0 MJ/kg (poultry), 0.52 MJ/kg (pig), and 0.52–0.53 MJ/kg (cattle). Loshakova et al. [34] evaluated grain quality of F₈–F₁₀ lines originating from crosses Trititrigia × Elytrigia intermedia and Trititrigia × Elymus fractus. These crosses were made to create new sources of valuable traits for the breeding of bread wheat and as “bridges” for further transfer of the traits in breeding. The study showed that both studied combinations had high grain quality. According to the sedimentation indices of the combination Trititrigia × E. intermedia, 4 samples out of 15 were classified as valuable wheat, while 11 samples belonged to the strong wheat quality group. Gluten content was high—from 30% to 35%. In the crossing combination Trititrigia × E. fractus, 3 samples belonged to valuable wheat and 12 samples were in a group of strong wheat.

Cultivar Pamyati Lyubimovoy (in memory of Lyubimova, N.V. Tsitsin’s colleague and friend) was the first Trititrigia officially released in Russia in 2020. The cultivar originated from a cross between bread wheat Istrinka with Agropyron glaucum and A. elongatum and further crossed with Trititrigia [31]. The grain yield was in a range of 3 t/ha, wet biomass—55 t/ha, and hay—14 t/ha. Grain weight (1000 kernels) varied from 27 to 32 g. High resistance to leaf rust and excellent regrowth capacity after harvesting for grain was demonstrated. Spike threshing is slightly tighter compared to bread wheat [35], requiring combine harvester special adjustment.

Pamyati Lyubimovoy was extensively studied in the fertile Rostov region [36] in 2021 and 2022. It was found that the average biological yield of Trititrigia in the southern zone of the Rostov region in two years was 4.28 t/ha, which was 1.57 t/ha less than control winter wheat Stanichnaya. The weight of the straw of Trititrigia was 1.9 times higher than control. Technological indicators of the quality of Trititrigia grain corresponded to the first quality class for protein (more than 19%), gluten (33.3%), and falling number (274 s); the third class according to the gluten deformation index (GDI) (81.5 points); the fifth class according to test weight (691 g/L). The general baking evaluation of Trititrigia grain allowed it to be classified as valuable wheat (Figure 3). However, Rostov conditions of hot and dry summers did not allow any regrowth of Trititrigia after harvest and its potential was not realized [37].

In summary, the breeding and research work at the Main Botanical Garden initiated by N.V. Tsitsin has continued with the preservation of his genetic resources, new targeted wide crosses programs, germplasm screening, and evaluation. New methods and tools have been utilized. The main focus presently is on regrowing types of material and much less on truly perennial wheat. Introgression of genes from wild species to wheat has successfully contributed to new cultivars released and grown in the country.

5. Perennial Wheat in Siberia

Perennial wheat work was initiated by N.V. Tsitsin in Omsk in 1932, and research resumed in 2016 when an international collection with 25 accessions of perennial cereals was first evaluated at Omsk State Agrarian University as part of a global trial. The results of evaluation of “early generation” winter perennial cereals were summarized by Hayes et al. [15]. Only six genotypes survived the first winter and exhibited valuable agronomic traits: CPI147235a (T. aestivum × Th. ponticum × T. aestivum) and 11955 (T. aestivum × Th. ponticum), both from USA, Washington State University; Agrotana (T. aestivum × Th. ponticum, USA); OK7211542 (T. aestivum × Th. ponticum, USA, Oklahoma State University); TAF46 (T. aestivum × Th. intermedium, France); and Otrastayushchaya 38 (T. aestivum × Th. intermedium, Russia). However, none of the wheats survived the second winter and did not demonstrate the persistence required for perennials.

Study and evaluation of six selected perennial wheats continued but as annual winter wheat suitable for harsh Siberian conditions during 2018–2020 [38,39]. The genotypes CPI147235a and Otrastayushchaya 38, obtained on the basis of Th. ponticum and Th. intermedium, were distinguished by high winter survival with averages of 78% and 90%, respectively, over 2019–2020. Depending on the weather conditions of the year, accessions 11955, TAF46, and Otrastayushchaya 38 formed acceptable grain yields of 240–290 g/m² over three years [38]. The collection accessions were characterized by the ability to regrow after harvest (Figure 4). Perennial wheat germplasm demonstrated high content of protein and gluten in grain. On average for four years of research, protein content varied from 19.3% (CPI147235a) to 21.2% (Otrastayushchaya 38), which surpassed control winter wheat Omskaya 4 (14.7%). The gluten content was 41.0–47.6%, which was significantly higher compared to Omskaya 4 (28.1%) [39].

Selected perennial wheat genetic resources were crossed with winter wheat varieties from Russia, USA, and International Turkey-CIMMYT-ICARDA program to expand genetic diversity and utilize valuable traits in winter wheat breeding. A number of constant lines originating from these crosses are being tested in field trials and demonstrated good winter survival, with grain yields varying from 150 to 470 g/m² in 2024.

6. Intermediate Wheatgrass Genetic Resources as Perennial Grain

Intermediate wheatgrass (IWG) (Thinopyrum intermedium = Agropyron intermedium = Agropyron glaucum) is a species native to the Eurasian continent and widely grown across steppe, forest–steppe, and desert regions [40] (Figure 5). This is a forage crop in Russia with seven cultivars officially registered for use nationwide [23]. However, this crop is considered a non-conventional forage, with a limited area of production [41].

One of the recent IWG forage cultivars, Ufimets, was developed in Central Russia by cross-pollination of locally collected forms with variety Rostovskiy 31 followed by several cycles of selection [42]. Vegetation period from spring regrowth to heading (mowing ripeness) is 55–67 days and 98–105 days to full maturity. The cultivar has exceptionally high winter hardiness, drought resistance, and disease resistance. The seed productivity of Ufimets averaged at 0.65 t/ha over 6 years with a 1000-kernel weight of 6.5 g. The yield of green biomass for 2 mowings was 33.4 t/ha. The optimal technology for this cultivar production includes early spring planting with inter-row spacing of 60 cm and seeding rate of 8 kg/ha.

Substantial research on IWG biology and genetics was conducted at the Institute of Cytology and Genetics in Novosibirsk, Western Siberia [43]. The work on the gene pool of wheatgrass began in 1971 when 90 collected plants were allowed to pollinate freely, and the seeds from this population formed a basic nursery of 1000 plants (I₀), 5–15 from each of the parent forms. Then, all plants were subjected to forced self-pollination. By 1983, a collection of more than 2000 plants of various degrees of inbreeding (I₀–I₃) had been created. Great diversity between plants in the weight of 1000 grains and their color, height, and width of leaves was revealed. In further works, partially homozygous plants of wheatgrass up to the 5th generation of inbreeding were obtained by forced self-pollination. The best of these plants had been used in crosses with winter wheat varieties.

To accelerate the breeding process, homozygous lines of wheatgrass were developed through anther culture, although less than 1% of genotypes had the ability to promote the haploproduction of green androgenic plants [44]. A. glaucum haploids exhibited very high frost resistance, with the freezing time before loss of turgor in leaf cells 30–70 times higher than the most frost-resistant winter wheat. After vernalization, haploid plants were planted in the field in early spring. In the first two years, haploids, as usual for perennial plants, gradually increased the mass of the root system and productive tillering. At the same time, the height of plants, as well as the size of the leaf, ear, and anthers, were significantly reduced in comparison with the parent diploid plants. After 3 years, some of the plants significantly increased their habit and acquired the characteristics of parent diploid plants. Cytological analysis showed the presence of a diploid set of chromosomes (2n = 42) in these plants, showing spontaneous duplication of chromosomal set. For the period from 1987 to 2008, a genetic collection (more than 500 plants) of haploids, doubled haploids, and donor plants of wheatgrass was developed.

The work on IWG as a forage crop or as a parent for wide crosses with wheat never resulted in breeding it as a perennial grain crop. The pioneering work of the Land Institute to domesticate IWG by removing unfavorable traits and increasing seed size signified a step forward in perennial path vision, which was not used in the species’ native land. Recent research on the genetic origin of food-grade IWG is focused between the Black Sea and Caspian Sea in the Stavropol region of Russia, with smaller contributions likely from collections as distant as Kazakhstan in the east to Turkey in the west [45]. It was natural that some early IWG populations from the Land Institute were shared with CIMMYT and with Omsk State Agrarian University to initiate breeding programs for perennial grains.

IWG cultivar Sova was developed and officially released by Omsk State Agrarian University for Siberia and its neighboring regions [46]. The initial population was obtained from the Land Institute and improved for adaptation to local conditions and productivity [47]. The yield of grain, green biomass, and hay in Sova variety during the three years averaged at 0.92, 21.0, and 7.1 t/ha, respectively (Figure 6). Grain quality indicators were high—19.4% protein and 36.3% gluten content. The average number of grains in the spike was more than 50, and 1000-kernel weight was 9.7 g. The length of the roots of the Sova variety was 6.9–9.8 times greater, and the surface area of all roots was 8.0 times greater than that of winter and spring bread wheat. The total number of agronomically important groups of microorganisms in IWG Sova root horizon turned out to be 2.2 times higher. The cultivar is demanded by farmers and primarily used as a forage crop since the grain is not marketed yet. The estimated production area in 2024 was around 5000 ha. The new IWG cultivar Filin from Omsk State Agrarian University, officially released in 2024, is characterized by high winter hardiness, drought tolerance, and purple color of grain with high content of protein (19.7%) and gluten (38.1%). The cultivar Filin can grow in one place for 7 years and is recommended both for green biomass and grain.

A study by Aydarov et al. [38] evaluated genetic gains in IWG populations for plant height and 1000-grain weight under Omsk conditions. Clones were selected according to plant height and other traits for population improvement. Plant height gains were preserved in subsequent generations. The selection by 1000-grain weight increased the value of this parameter by 30%. The coefficient of genetic determination for this trait was 0.96, indicating a high contribution of the genotype and the effectiveness of genetic gain. Selection for plant height affected the spike length and weight, number of grains and their weight per spike, as well as 1000-grain weight. Superior populations with large grains exceeding 12.5 g were identified.

The potential of Sova grain as a novel ingredient in breadmaking was extensively studied in Omsk in collaboration with Turkish institutions [9]. Enrichment of regular flour with IWG flours significantly decreased Zeleny sedimentation and gluten index values and increased the dry and wet gluten contents. The bread’s yellow pigment content and crumb b* color value increased with the increasing level of IWG supplementation. IWG addition also had a positive effect on phenolic and antioxidant properties. Bread with 15% IWG substitution had the highest bread volume (485 mL) and lowest firmness values (654 g–force; g–f) compared to the other breads, including the control (i.e., wheat flour bread). The results indicated that IWG has great potential to be used in bread production as a novel, healthy, and sustainable ingredient. Trevisan et al. [48] evaluated the pasting and rheological properties of doughs made from bread wheat flour and blended bread wheat–IWG flours (IWG content: 15%, 30%, 45%, and 60%). The RVA peak, trough, and final viscosity values significantly (p < 0.05) decreased, and the pasting temperature increased as the IWG substitution level increased. IWG-containing flour blends had a lower retrogradation tendency compared to the control. As the IWG flour substitution level increased, both empirical (Mixograph, Kieffer dough and gluten extensibility, Glutograph) and fundamental (linear oscillatory frequency sweep and creep–recovery) rheological measurements indicated relatively weaker dough properties imparted by the addition of the IWG flour. Obviously, IWG bread-making grain quality improvement is needed for bread production.

7. Perennial Rye and Triticale

N.V. Tsitsin inspired a number of scientists to work on perennial grains (Goncharov, 2023) [6]. A.I. Derzhavin was a remarkable breeder who made great contributions to wide crosses and development of perennial grains [49]. Born in the Penza region, he graduated from Voronezh Agricultural Institute and his main work was conducted in the Stavropol region of South Russia. He started to work on perennial rye in the 1930s by crossing Secale cereale (cultivar Tarashanskaya) × S. montanum [50]. The resulting crop was described as the subspecies of rye S. cereale ssp. derzhavinii (Tzvelev). Perennial rye cultivar Derzhavinskaya 29 was officially registered in 1981 and is still included in the State Register [23]. This rye is a diploid with 2n = 14.

The main use of perennial rye was forage due to its ability to produce biomass and regrow after mowing. The crop was well studied across Russia. Shebarskowa and Kipaeva [51], using second- and third-year crops, obtained 28.5 and 13.6 t/ha of wet and dry biomass, respectively, with 2 t/ha grain yield in the low Volga region. The crop tolerated very well additional biomass harvest and grazing. In 2005–2012, field experiments studied forage productivity of perennial rye Derzhavinskaya 29 in both its pure form and mixed with sainfoin in comparison with winter rye and winter triticale in the Ural Mountains [52]. Perennial rye had high winter hardiness and drought tolerance with the period of economic use of 2–3 years. After overwintering, when harvesting green mass for forage, it was able to give another harvest in July, which is about 35% of the total biomass.

Perennial rye is recommended to be sown in early spring under the cover of grain crops, as well as at the end of summer in its pure form as a winter crop [52]. A joint sowing of perennial rye with sainfoin under the cover of oats in spring was proposed. In the year of sowing, oats mixed with sainfoin were harvested for forage. In the second year, a mixture of perennial rye with sainfoin was mowed for feed; in the third year, perennial rye was harvested for grain (yield of 0.8–1.5 t/ha) and biomass. Bektyashkin [53] studied the effect of five seeding dates (spring to late summer and early fall) and six seeding rates (3, 3.5, 4, 4.5, 5, 5.5 million seeds per ha) on perennial rye grain and biomass yield in the middle Volga region. The highest average grain yield, averaging for 2nd, 3rd, and 4th seasons (2.1 t/ha), was obtained upon planting on August 15 with 4–5.5 million seeds/ha. In Western Siberia (Tyumen region), Derzhavinskaya 28 demonstrated yield of 2.4 t/ha, almost half that of regular rye varieties, with 1000-kernel weight of 27–30 g [54].

Derzhavin perennial rye was later transformed into tetraploid by Tsitsin and his colleagues at MBG in Moscow, and two cultivars were released (Snegirevsksya 28 and Utro) [6]. However, the tetraploid version of perennial rye was late in development and had lower winter hardiness, with few plants surviving the second winter [54]. Several other attempts at developing perennial rye were undertaken using crosses between different forms of S. montanum (cultivars Kormovaya 61 and GAK) (Shcheglov, 2009) [55]; variety R-208 was developed for irrigation with grain yield 10–20% higher compared to Derzhavin rye [56].

Perennial rye also proved its viability in similar environments in Alberta, Canada [57]. The material, developed through crosses between Secale cereale and S. montanum, was tested across several sites and demonstrated superior biomass compared to winter and spring rye during the two seasons, though the grain yield was almost 50% of that of the regular rye.

The perspectives of perennial rye have been supported by a long-term experiment of permanent cultivation of annual winter rye for 126 years as reported by Belyavsky [58]. The yield of winter rye for this period averaged at 1.21 t/ha. The control was the usual sowing of winter rye in the crop rotation, with the grain yield at a level of 2.5–3.5 t/ha. The most common insects were thrips, cereal aphids, grass bugs, and cereal fleas, causing only local damage in some favorable years. Among the diseases, the most frequently observed were root rot (Bipolaris sorokiniana Shoem.), brown rust (Puccinia recondita Rob. et Desm.), septoria (Septoria tritici Rob. et Desm.), and tan spot (Drechslera tritici-repentis It). The long-term phytosanitary situation of perennial rye stabilized and did not cause crop failure. Thus, there is a likelihood that perennial rye being grown in only a few years would not be substantially affected by pests and diseases.

A.I. Derzhavin made efforts to develop perennial triticale [50]. The first hexaploid wheat–rye amphidiploid (2n = 6x = 42) was synthesized by him in 1932 as a result of hybridization of semi-winter durum wheat Leukurum 1364-1 with wild perennial rye S. montanum from Armenia. The first hybrid generation was sterile. Repeated pollination of F₁ hybrids with the same rye resulted in a fertile plant that gave rise to a perennial hexaploid amphidiploid [59]. In the 1960s and 1970s, Derzhavin used complex crosses {(Triticum aestivum cv. Alabasskaya × Secale derzhavinii) × [(T. aestivum cv. Leucurum 1364/1 × perennial wild rye) × S. derzhavinii]} × (T. durum × A. glaucum) to develop perennial triticale. The best results in producing perennial triticale were obtained by selection for regrowth of winter-type tillers and for a vigorous root system while hybridizing perennial triticale with perennial wheat–rye–Agropyron hybrids. Presently, no references are available on this genetic resource and its fate is not clear. Derzhavinskaya 28 perennial rye and its derivatives can contribute to international efforts in breeding perennial grains.

8. Perennial Sorghum

A.I. Derzhavin also implemented the idea of developing perennial sorghum (Sorghum derzhavinii Tzvel.) by crossing Sorghum bicolor with Sorghum halepense [60]. Two perennial sorghum cultivars, Karavan and Travinka, were officially released in 2004 and are included in the State Register [23]. Perennial sorghum’s main purpose was forage production.

One area where perennial sorghum has been studied and used is North Pre-Caspian arid pastures with 150–250 mm of annual precipitation. Derzhavin sorghum was targeted for the reclamation of degraded pasture ecosystems and creation of intensive, high-nutrient hayfields [61]. Two varieties of perennial sorghum—Travinka and Karavan—were grown with different planting densities: 10,000, 20,000, and 40,000 plants per hectare under irrigation and dryland. Natural pasture was the control. The highest green biomass yield was observed in the variety Travinka at a density of 40,000 plants per hectare—41.6 t/ha under irrigation and 7.9 t/ha under rain-fed conditions; in the variety Karavan, density of 20,000 plants per hectare was optimal and produced 24.8 t/ha and 3.9 t/ha of biomass, respectively. Analysis of the chemical composition showed that, in comparison with the natural pasture, sorghum was more nutritious.

A study in the Chechen Republic also demonstrated superior drought tolerance and forage quality performance of perennial sorghum, with a green biomass yield of 4–8 t/ha [62]. Sugar content in perennial sorghum exceeded 80 g/kg, well above most forage crops. Perennial sorghum Travinka demonstrated higher water-use efficiency compared to sugar sorghum and sorghum–Sudan grass hybrids in the middle Volga region (Tatarstan Republic) [63]. Unfortunately, the references lack information on the perennity of Derzhavin sorghum and the grain yield. Nonetheless, the perennial sorghum genetic resources are viable and available for production and further improvement. Russian perennial sorghum can be a valuable genetic resource to contribute to existing efforts in breeding this crop in Africa and other continents.

9. Summary and Perspectives

Nikolay Tsitsin, a pioneering Russian scientist, first proposed perennial wheat as an alternative crop in the early 1930s by crossing wheat with perennial grasses. His work led to creating stable wheat amphiploids with some perennity and successful gene transfers from wild wheat relatives to cultivated varieties. Tsitsin’s work resulted in two main groups of genetic resources: perennial wheat (which never reached commercial cultivation) and regrowing wheat cultivars that were officially registered. His most practical outcomes were wheat–wheatgrass hybrids developed through backcrosses to wheat, with six spring wheat varieties widely cultivated in Western Siberia and Northern Kazakhstan in the 1960s. The breeding work at the Main Botanical Garden continued after Tsitsin, maintaining a collection of over 250 forms of Trititrigia developed through complex crosses. Tsitsin’s original wheat–wheatgrass hybrid lines have been preserved at MBG. Part of the collection is stored at the Vavilov Institute in St. Petersburg. The germplasm is accessible subject to the policy of the holding institutions.

Intermediate wheatgrass is native to Eurasia and grown across Russia mainly as forage. In 2022, Omsk State Agrarian University released the IWG cultivar Sova for grain production, derived from the Land Institute populations but improved for local adaptation. It produced about 0.9 t/ha grain yield and 21 t/ha green biomass. Studies on its potential as a novel baking ingredient showed promising results.

A.I. Derzhavin developed perennial rye (Secale cereale ssp. derzhavinii) in the 1930s by crossing cultivated rye with wild perennial rye. Cultivar Derzhavinskaya 29 was officially registered in 1981 and remains in the State Register. It is primarily used for forage production due to its regrowth ability. A.I. Derzhavin also developed perennial sorghum (Sorghum derzhavinii) by crossing cultivated sorghum with wild perennial Johnson grass. Two cultivars, Karavan and Travinka, were released in 2004. They have shown superior drought tolerance and forage quality in the North Pre-Caspian arid regions with very high biomass production.

The main challenges for perennial grains in Russia are development of modern varieties suitable for existing cropping systems and development of relevant production technology. In breeding, it is important to exchange germplasm and genetic resources between institutions in Russia and with advanced foreign programs. Application of modern physiological tools, precision phenotyping, and genomic methods would contribute to the rapid development of competitive cultivars. Once new cultivars are available, production technologies ought to be developed for optimal use of resources and profitable grain production. As the concept of perennial grains develops, breeding and agronomy research ought to be reoriented toward grain and perennity, focusing largely on its potential for use across the country.