Spanish Spelt Wheat: History
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Subjects: Agronomy
Contributor:
Juan B. Alvarez

Spelt wheat (Triticum aestivum L. ssp. spelta Thell.) is an ancient wheat that was widely cultivated in the past. This species derived from a cross between emmer wheat (T. turgidum spp. dicoccum Schrank em. Thell.) and Aegilops tauschii Coss. Its main origin was in the Fertile Crescent (Near East), with a secondary center of origin in Europe due to a second hybridization event between emmer and hexaploid wheat. This species has been neglected in most of Europe; however, the desire for more natural foods has driven a revival in interest. Iberian spelt is classified as a geographical group differing to the rest of European spelt.

  • agrobiodiversity
  • ancient wheat
  • genetic resources
  • neglected crops

1. Introduction

The area known as the Fertile Crescent (Near East) has been widely suggested by numerous authors as the origin of wheat [1]. Under the name of wheat is possible to find numerous species and subspecies from the genus Triticum, grouped in three ploidy levels: diploid (2n = 2× = 14, AmAm), tetraploid (2n = 4× = 28, AuAuBB) and hexaploid (2n = 6× = 42, AuAuBBDD). Wild ancestors have been found for the diploid and tetraploid species, which were domesticated by hunter-gathering societies during their evolution to farming societies [2]. The first archaeological traces of wheat have been identified as from around 10,000 BP (pre-pottery Neolithic A), being mainly diploid and tetraploid species. In contrast, all hexaploid wheats are domesticated, having their origin in natural crossing between emmer wheat (Triticum turgidum spp. dicoccum Schrank em. Thell., 2n = 4× = 28, AuAuBB), the first domesticated tetraploid wheat, and a wild grass (Aegilops tauschii ssp. strangulata Coss., 2n = 2× = 14, DD), which could have at least two separate origins: one in Iran and another in Turkey or northern Syria [3]. This event was recreated by McFadden and Sears [4], obtaining plants with high similitude to the ancient wheat known as spelt (T. aestivum L. ssp. spelta Thell., 2n = 6× = 42, AuAuBBDD), and considered to be the ancestor of common wheat (T. aestivum L. ssp. aestivum, 2n = 6× = 42, AuAuBBDD).
A third origin of spelt outside the Fertile Crescent has also been suggested [5]; European spelt could be derived from a secondary hybridization between emmer wheat and hexaploid wheat, probably club wheat (T. aestivum L. ssp. compactum Host em. Mackey) [6][7][8][9]. In addition to this, European spelt could be classified into two eco-geographical groups: Iberian (pol. ibericum Flaskb.) and Bavarian (pol. bavaricum Vav.) groups, being the first originated from Asia and show great differences to the spelt from Central Europe that likely originated following secondary hybridization [9][10]
Some morphological characteristics of spelt clearly differ from common wheat, including non-free threshability, brittle rachis, long spikes and hulled grains [11][12][13][14][15], which mainly arose from the expression of two loci: Q and Tg genes. The Q gene encodes a transcription factor (APTETALA2 family) related to these characteristics [16][17]. The first studies found the presence of a recessive allele (q) in common wheat that is key in the morphological differentiation between spelt and common wheat. This locus is on the long arm of chromosome 5A in polyploid wheat [6][18]. The other gene, Tg, is related to glume tenacity (i.e., hulled grain) and located on the short arm of chromosome 2 of each wheat subgenome. The first gene evaluated was Tg-D1, found by Kerber and Rowland [19], in populations derived from crosses between durum wheat and Ae. tauschii, and the observed non-free-threshing trait was due to the presence of a dominant allele (Tg-D1) from Ae. tauschii. This suggested that this gene was epistatic to the Q gene. Later, orthologous genes were detected in chromosome 2BS (Tg-B1) by Simonetti et al. [20], using crosses between durum and emmer wheat, and in chromosome 2AS (Tg-A1) by Dvorak et al. [21] in spelt. Data suggested that mutations of Tg-A1, Tg-B1 and Q present in emmer wheat were responsible for the free-threshing trait in modern durum wheat [22]. This process was later carried out in hexaploid wheat (spelt), including the mutation on Tg-D1 (Figure 1).
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Figure 1. Evolution of genes related to the domestication in Triticum species. The free-threshing character is controlled by the Q gene and the tenacious glume by the Tg gene.
In recent times, the presence of these allelic variants in any modern material has been sufficient for considering as spelt. This has led to two different spelt types: traditional/pure and modern/synthetic [23]. The latter are derived from crosses between semi-dwarf common wheat and traditional spelt, which show spikes with speltoid morphology and tenacious glumes. In contrast, traditional spelts are obtained by selection among old spelt landraces. Although both options are completely feasible, the differences between them should be made clear for free choices by consumers.

2. Recovery or Development of New Spelt Varieties

Parallel with the return to more natural food, the interest in traditional products has clearly increased. However, in many cases, these products are currently made with modern flours, and so some properties associated with these traditional products differ. For this reason, the renewed interest in these products has increased the need to recover the old materials used to make them in the past, together with investigation of the quality characteristics of these old materials to design new materials with similar characteristics.
Faced with these circumstances, two ways have been used with more or less success: use of old landraces vs. generating new varieties by crosses with common wheat. The main way is evaluating old landraces and identifying the best lines. This process implies that phenotypic selection within landraces, together with correction of undesirable traits (mainly the lodging due to the high plant height), has been carried out by cultural techniques: low fertilization or increasing seeding density. Spelt has been grown in some European regions with traditions in the maintenance of spelt until recent times. Some of these zones have designed protection mechanisms for these ancestral products regulated by European Union laws, such as the Protected Geographical Indication (PGI), which is key to the maintenance of pure spelt. One example of this is the PGI «Farro della Garfagnana» (Italy) [24]. One similar process has been valued in several occasions for the Asturian spelt (escanda asturiana), although without positive results until moment.
Our team developed one set of 32 lines using this way, which were obtained from different Spanish landraces included in the 1930s collection, and selected according with the HMWGs composition [25]. For this, one unique seed was used to generate each line, removing any possible heterogeneity present in the landraces used (Figure 6). The additional selection among these materials has allowed the identification of one promising line with good adaptation to the climatic and edaphic conditions of Asturias (unpublished results). This tendency to local adaptation is frequent in traditional materials, and makes selection difficult in other environments.
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Figure 6. Recuperation of old landraces in Asturias (Northern Spain).
The second way to develop spelt varieties is generating hybrids by crosses with common wheat (Figure 7) [26][27]. For the development of lines from these hybrids, the offsprings should be self-pollinated during several genenerations, selecting those plants that, additionally to good agronomical traits, present spikes with speltoid morphology (allele q) and tenacious glumes (allele Tg) (Figure 1). This process, in general long in the time, can be shorted using speed breeding techniques [28]. Alternatively, the anther culture for doubled haploid production can be used with this same finality [29][30][31].
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Figure 7. Spikes of spelt × common wheat hybrids found in Asturias during the collecting mission of 2004.
The main objective of the new spelt varieties is the correction of some agronomical problems such as lodging by producing shorter plants, which permit an increase in harvest index [32]. These materials have been obtained by hybridization with semi-dwarf common wheat, and European cultivars as Alkor, Badengold, Ceralio and Cosmos are good examples. In any case, the resulting varieties should maintain morphologic characteristics similar to traditional spelt, including spike morphology and tenacious glumes, because without these specific characteristics they cannot be considered to be spelt. In any case, the studies carried out with these new spelt varieties showed that, although their grain yields are clearly higher than those traditional spelt, these are sensitive lower than those of common wheat [33][34][35].
There has so far been little development of new spelt varieties (deriving from crosses with common wheat) in Spain or with Spanish germplasm, with two cultivars being registered: Annamaria and Viso. However, this could be important in coming years due to the renewed interest in this crop. In fact, new materials are being developed using both the double-haploid production [36] and the self-pollination and selection by speed breeding (unpublished results). Nevertheless, the likelihood of genetic erosion in the traditional material due to the presence of these new materials should be considered. Obviously, agriculture is an economic activity, and the loss of profitability could motivate some traditional growers to neglect the old material for more modern spelt that, a priori, could be more profitable due to lower crop losses. In this case, traditional spelt would only be cultivated by the few aged farmers who continue to use it for home consumption and could be permanently lost when they pass.

3. Perspectives

In recent times, spelt has become a fashionable crop in some areas of Europe, including Spain. Although this species was widely cultivated in the past in Spain, during the 20th century it suffered deep erosion that led it to near extinction [37]. This deep erosion caused that the available material was mostly stored in some germplasm banks, being scarce in the fields [38].
The revival of spelt cultivation at recent decades did mainly stock up from the scarce material available by farmers, which implied a very narrow genetic base. For this reason, it is necessary to evaluate and recover materials from germplasm banks, which have not been accessible for pioneer farmers in the recovery of spelt as a crop.
Nowadays, similar to what has been performed in other parts of Europe [39][40][41][42][43][44]; spelt improvement programs have been developed in Spain. Although some of them have been carried out by the selection within traditional materials [45][46], others have used the hybridization with semi-dwarf common wheat in order to obtain spelt varieties with characteristics agronomic more suitable to current agricultural techniques [36]. These new varieties present low-height plants that allow their mechanization and avoid the habitual lodging in traditional materials. Nevertheless, regardless of the fact that the growing consumer demand has led to a gradual increase in the cultivated area, it still remains at low levels, so there are no official statistics in this regard.
This trend must be linked to conservation programs of the genetic resources of the species, to avoid possible erosion and to develop higher variation levels in the commercialized materials. Furthermore, the studies carried out to assess the variability levels of the materials deposited in germplasm banks have shown not only the uniqueness of the material from the Iberian Peninsula [10], but also the presence of some variation levels for, among others, technological quality characteristics [47][48][25][49][50][51][52][46][53][54][55], which may allow an important diversity within the varieties of spelt available.
Due to the current trend to seek additional nutritional characteristics in crops, the development of spelt varieties with nutraceutical characteristics due to the presence of trace elements, can be a good tool for the maintenance and expansion of this crop both in Spain and elsewhere of the world.

This entry is adapted from the peer-reviewed paper 10.3390/plants10122748

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