Wild Relatives of Cultivated Plants in India. A Reservoir of Alternative Genetic Resources and More,A.K. Singh,Springer (2017) 309 pp., Price 156.99 £, ISBN: 9789811051166

Wild relatives of cultivated plants are the treasure trove of genetic resources for crop improvement and dietary diversification. Especially, they are crucial for conferring diseases resistance and abiotic stress tolerance (such as drought, flood, heat-shock, salinity, etc.) in cultivated crops and thereby breeding next generation of climate smart crops (i.e. climate resilient crops) for futuristic climatic conditions. Unfortunately, most of the wild relatives of crop plants across the world are being neglected and underutilized, without recognizing its real potential. In this backdrop, the book herein review titled “Wild Relatives of Cultivated Plants in India: A Reservoir of Alternative Genetic Resources and More”, authored by Anurudh Kumar Singh, National Bureau of Plant Genetic Resources, New Delhi, India is a timely and topical endeavor to provide a vivid account of more than 958 species of wild relatives of cultivated plants in India along with their occurrence, distribution, current status, as well as their future potential for breeding program and dietary diversification. Moreover, the author has also explored the additional utility of such crop species for ecosystem restoration, phytoremediation and soil carbon sequestration in detail. Here we distill the essence of this 309-page book for the better comprehension and understanding for a wide range of potential readers like graduate students and doctoral researchers in agricultural and environmental sciences, agronomist, plant breeders, as well as various national and international policy makers and regulatory agencies.     

Genetic diversity and relationships between wild and cultivated olives (<Emphasis Type="Italic">Olea europaea</Emphasis> L.) in Sardinia as assessed by SSR markers

The genetic relationships within and between wild and cultivated olives were examined and clarified in an isolated and restricted area, such as the Mediterranean island of Sardinia. Wild (21 individuals) and cultivated olive trees (22 local cultivars from a germplasm collection and 35 ancient trees) were genotyped by means of 13 SSR loci. Five cases of synonymy were observed and nine distinct genotypes were identified in the collection. Five novel genotypes were also detected among the ancient trees. Differences on the allelic composition and heterozygosity levels were found between wild and cultivated trees. Model-based clustering method classified the olive trees into two major gene pools: (a) wild genotypes and (b) local cultivars from the collection and from heritage olives. Regarding the cultivated plant material, we observed that: (a) most of the Sardinian cultivars shared the same allelic profiles with the ancient cultivated trees and (b) the majority of these cultivars and all the novel genotypes were not related to any other cultivars included in this study. These findings as well as the detection of unique alleles and a certain wild genetic background at some cultivars revealed by the Bayesian analysis may indicate their autochthonous origin. The synonymy cases found between local cultivars and Italian mainland cultivars indicate interchange of genetic material among these growing areas, suggesting thus a possible allochthonous origin. The information obtained can assist in the management of an olive collection and sheds some light on the survival of true oleasters and the origin of Sardinian cultivars.     

Spontaneous gene flow and population structure in wild and cultivated chicory, Cichorium intybus L.

Spontaneous gene flow between wild and cultivated chicory, Cichorium intybus L., may have implications for the genetic structure and evolution of populations and varieties. One aspect of this crop-wild gene flow is the dispersal of transgenes from genetically modified varieties, e.g. gene flow from GM chicory to natural chicory could have unwanted consequences. With the purpose to identify and quantify crop-wild gene flow in chicory, we analysed introgression in 19 wild chicory populations and 16 accessions of chicory varieties and landraces distributed across Northern, Central and Mediterranean Europe. The analysis used 281 AFLP markers and 75 SSAP markers giving a total of 356 polymorphic markers. Results from model based assignments with the program STRUCTURE indicated many incidents of recent gene flow. Gene flow was observed both between cultivars and wild populations, between landraces and wild populations, between different wild populations as well as between cultivars. Population structure visualized by distance-based clustering showed a North–South geographical structuring of the wild populations, and a general grouping of the cultivars corresponding to known origin. The results indicated, however, that the structuring between the two groups of wild and cultivated types was weak. As crop and wild recipients are genetically close and genes are transferred between the two types rather frequently, focus on mitigating crop-wild gene flow should be increased, before transgenic varieties are cultivated openly.     

Natural hybrids between cultivated and wild sunflowers (<Emphasis Type="Italic">Helianthus</Emphasis> spp.) in Argentina

Two introduced wild species Helianthus annuus L. and H. petiolaris Nutt. have become widespread in central Argentina and overlap the sunflower crop region. Intermediate off-type plants between the wild and cultivated species are often found, which is of concern because of the recent release of imidazolinone resistant varieties and the likely use of genetically modified sunflower cultivars. The progeny of 33 off-type plants obtained from 14 representative sites of the diffusion area were studied to confirm hybrid origin. Germination, survival, morphological traits and days to flowering confirmed hybridization between crop and both wild species, when compared to eight accessions of typical wild plants. Some progenies were presumably crop–wild H. annuus hybrids, some originated from the cross of cultivated plants and H. petiolaris, and two were the advanced generation of a cultivated hybrid. Hence, morphological traits are a good clue for the identification of spontaneous hybrid plants at field. The results indicate that crop–wild hybridization and introgression occur at various places in central Argentina. This fact may represent a way to herbicide resistance escape and future transgene escape if GM sunflower cultivars are released for commercial use.     

Utilization of the genetic resources of wild species to create a nontransgenic high flavonoid tomato

Flavonoids represent a large and important group of plant natural products that are ubiquitous in the plant kingdom. Epidemiological studies have shown the health benefits of a diet high in flavonoids. However, the dietary intake of flavonoids in most western populations is limited, creating a need to find alternative food sources for these polyphenolic secondary metabolites. The domestication of many of our cultivated food crops has resulted in alterations in the biosynthetic pathways of many essential micronutrients and vitamins through inadvertent counterselection against nutritional traits in favor of agronomic ones. Flavonoids are nearly absent from fruits of cultivated tomato (Lycopersicon esculentum Mill.), a major vegetable in human diets. Previous attempts to restore the flavonoid pathway in tomato fruits have been limited to transgenic strategies, suggesting that the problem was intractable through traditional methods. Here, we describe for the first time a nontransgenic metabolic engineering approach to developing a high flavonoid tomato using a wild tomato species (Lycopersicon pennelliiv. puberulum) and demonstrate the opportunities for restoring functional pathways using the genetic resources of wild species, resulting in production of healthier foods.     

Close Genetic Proximity Between Cultivated and Wild Bactris gasipaes Kunth Revealed by Microsatellite Markers in Western Ecuador

Bactris gasipaes Kunth (peach palm or Pejibaye) is the only domesticated palm of the Neotropics. The genetic relationships between the crop and its wild relatives are still unclear. We undertook field and laboratory work in order to describe differentiation and relationships between the wild and cultivated populations of the species in Western Ecuador, and their possible interactions. A volumetric study was undertaken on the fruits of both populations, as well as a population genetic analysis in order to clarify these relationships. Fruits from cultivated plants collected in the region of sympatry of wild and cultivated plants in North-West Ecuador showed intermediate volumes between those of reference samples for the wild and cultivated plants in allopatry. Using 8 microsatellite loci, we assessed 83 wild and cultivated individuals from Western Ecuador and cultivated plants from Amazonia and Central America as a reference for the cultivated form. We detected high polymorphism in the wild and cultivated samples and low, but significant level of genetic differentiation between wild and cultivated populations. The cultivated population in North-Western Ecuador showed close genetic proximity with the sympatric wild population, consistent with the volumetric study. These results have implications for hypotheses on evolution of this crop and for strategies of genetic conservation of the wild forms.     

Population structure of the primary gene pool of Oryza sativa in Thailand

The gene pool of cultivated Asian rice consists of wild rice (Oryza rufipogon Griff.), cultivated rice (O. sativa L.) and a weedy form (O. sativa f. spontanea). All three components are widespread in Thailand, frequently co-occurring within fields and providing the opportunity for gene flow and introgression. The purpose to this study is to understand the on-going evolutionary processes that affect the gene pool of rice by analysis of microsatellite variation. Results indicate that O. rufipogon, the wild ancestor of rice, has high levels of genetic variation both within and among populations. Moreover, the variation is structured predominantly by annual and perennial life history. High levels of variation are detected among cultivars indicating Thai cultivated rice has a broad genetic base with only a 20 % reduction in diversity from its wild ancestor. The weedy rice populations reveal varying levels of genetic variation, from nearly as high as wild rice to near zero. Weedy rice is genetically structured into 2 groups. Some populations of invasive weedy rice are the result of hybridization and gene flow between local wild rice and local cultivated rice in the regions of co-occurrence. Other populations of weedy rice are genetically nearly identical to the local cultivated rice. The diversity analysis indicates that the rice gene pool in Thailand is a dynamic genetic system. Gene flow is ongoing among its three main components, first between cultivated and wild rice resulting in weedy rice. Weedy rice in turn crosses with both cultivated varieties and wild rice.     

Characterization of single nucleotide polymorphism in Tunisian grapevine genome and their potential for population genetics and evolutionary studies

In this study, two gene fragments corresponding to the VvMYBA1 and VvMYBA2 loci were sequenced on a sample of grapes including cultivated and wild accessions originating from Tunisia, Germany and France. A total of 42 SNPs were detected in the sequenced fragments giving an average of 1 SNP every 33 bp. High level of polymorphism was observed in the samples either in cultivated or wild accessions. Pattern of nucleotide diversity indicates a non departure from neutrality expectations for wild grapevine sample for gene VvMYBA1 and VvMYBA2 and for cultivated sample for gene VvMYBA1. However, a linkage to a selective sweep was revealed for cultivated grapevine gene pool in gene VvMYBA2. A genetic structure of the studied sample according to accession taxonomic status was revealed by the UPGMA clustering with a considerable overlap. This result was confirmed by significant but low genetic differentiation values between cultivated and wild sample. The number of migrants Nm based on sequence data information between Tunisian cultivars and Tunisian wild accessions showed a low level of gene flow between those germplasms. This finding indicates that Tunisian cultivars do not derive directly from local wild populations but could mostly correspond to imported materials introduced during historical times. However, the possibility that some cultivars derived from ancestral events of local domestication or cross hybridization with native wild plants was not completely excluded for Tunisian grapevine accessions.     

Potential renal acid load in the diet of children and adolescents: impact of food groups, age and time trends

OBJECTIVE: The impact of acid-base balance on health is widely accepted. Here, we describe the potential renal acid load (PRAL) in the diet of healthy German children and adolescents. DESIGN: The Dortmund Nutritional and Anthropometric Longitudinally Designed (DONALD) Study is an ongoing longitudinal (open cohort) study (start 1985) collecting detailed data on diet, growth, development and metabolism in infants, children and adolescents. SETTING: Research Institute of Child Nutrition, Dortmund. SUBJECTS: Seven hundred and twenty children and adolescents (351 boys and 369 girls), aged 3-18 years, provided 4187 yearly collected 3-day dietary records between 1995 and 2005. RESULTS: Mean daily PRAL was positive in all age/sex groups (6-21 mEq day-1), and significantly higher in boys than in girls after the age of 8 years, even when calculated as mEq MJ-1. Fruits, vegetables and potatoes had a negative impact on PRAL; cheese, dairy products, cereals/bread and meat/fish/eggs had a positive impact. In a mixed linear model, PRAL, expressed as mEq day-1 and mEq MJ-1, remained stable during the study period, since time trends of PRAL-relevant food groups countervail each other. PRAL intake (mEq MJ-1) was significantly positively associated (P < 0.0001) with fat intake (% of energy intake, %E), but negatively with carbohydrate intake (%E; P < 0.0001). CONCLUSIONS: The analysis of dietary habits in our sample of German children and adolescents showed a moderate excess of acidity. Especially older boys should be encouraged to eat more potatoes and vegetables as good sources of dietary alkalinity. The PRAL concept is compatible with current concepts for a healthy diet.     

The influence of European and American wild germplasm in hop (<Emphasis Type="Italic">Humulus lupulus</Emphasis> L.) cultivars

Microsatellite variation at the nuclear and chloroplast genomes was evaluated for wild European and wild American hops, in order to assess the genetic diversity and origin of cultivated hops. Seven nuclear loci and 32 chloroplast loci were used in the analysis of 182 hop accessions including wild European (68), wild American (48), and cultivars (66). A total of 116 alleles were identified using 7 nuclear microsatellites showing different averages of polymorphism and distribution in the wild American and European accessions and cultivars. Two main groups were established as revealed by several statistical analyses; one including European wild accessions and cultivars and a second group consisting of American wild accessions. Three polymorphic chloroplast microsatellite loci were detected, six alleles were scored which defined a total of five haplotypes that were exclusive or presented different distribution between American and European wild accessions. A major influence of the wild European haplotypes was detected among hop cultivars. To the best of our knowledge, this is the first work reporting the use of chloroplast microsatellites in hops.     

Growth and physiological characterization of low nitrogen responses in Tibetan wild barley (Hordeum spontaneum) and cultivated barley (Hordeum vulgare)

Development of crop cultivars with high yield under low nitrogen (N) supply is a basic approach for the enhancement of agricultural sustainability. The previous studies showed that Tibetan wild barley shows wider genetic diversity in abiotic stress and poor fertility tolerance. In this study, four barley genotypes (two Tibetan wild and two cultivated), differing in N use efficiency (NUE), were characterized for their growth and physiological responses to low N stress. The genotypes ZD9 (cultivated) and XZ149 (wild) with high NUE performed better in terms of shoot dry weight (DW) and photosynthetic parameters under both low and normal N levels and had higher antioxidative enzyme activities, N concentration, and accumulation in both shoots and roots under low N stress. The current results showed the substantial difference among barley genotypes in low N tolerance and verified the significance of Tibetan wild barley in the genetic improvement of cultivated barley in NUE.     

Structure-activity relationship (SAR) studies of benzoxazinones, their degradation products, and analogues. Phytotoxicity on problematic weeds Avena fatua L. and Lolium rigidum Gaud

Avena fatua L. (wild oat) and Lolium rigidum Gaud. (rigid ryegrass) are highly problematic weeds affecting a wide variety of cereal crops worldwide. The fact that both of these weeds have developed resistance to several herbicide groups made them optimal candidates as target organisms for ongoing research about the potential application of allelochemicals and analogue compounds as natural herbicide models. Benzoxazinones, a family of natural allelochemicals present in corn, wheat, and rye, including 2,4-dihydroxy-(2H)-1,4-benzoxazin-3(4H)-one and 2,4-dihydroxy-7-methoxy-(2H)-1,4-benzoxazin-3(4H)-one, together with some degradation products, found in crop soils as well as in other systems, and some synthetic analogues of them were tested on wild oat and rigid ryegrass seeds; the results were statistically treated, and some structure-activity relationships, useful in further development of natural herbicide models, were elucidated. The most active compounds were the synthetic benzoxazinone 2-acetoxy-(2H)-1,4-benzoxazin-3(4H)-one and the degradation product 2-aminophenoxazin-3-one, with highly significant inhibition on the development of both weeds. The ecological role of these compounds is discussed by considering both degradability and phytotoxicity. The bioactivity of aminophenoxazines has been correlated by their aqueous solubility-lipophilicity predicted by means of computational methods.     

Genetic structure and differentiation among grapevines (<Emphasis Type="Italic">Vitis vinifera</Emphasis>) accessions from Maghreb region

Three gene pools representative of Vitis vinifera L. subsp. vinifera (=subsp. sativa Beger) growing in the Maghreb regions (North Africa) from Tunisia (44), Algeria (31) and Morocco (18) and 16 wild grape accessions (Vitis vinifera L. subsp. sylvestris (Gmelin) Beger) from Tunisia were analysed for genetic diversity and differentiation at twenty nuclear microsatellites markers distributed throughout the 19 grape chromosomes. 203 alleles with a mean number of 10.15 alleles per locus were observed in a total of 109 accessions. Genetic diversities were high in all populations with values ranging from 0.6775 (Moroccan cultivars) to 0.7254 (Tunisian cultivars). F _st pairwise values between cultivated grapevine populations were low but found to be significantly different from zero. High F _st pairwise values were shown between wild and cultivated compartments. Two parent offspring relationships, two synonyms and two clones of the same cultivar were detected. The rate of gene flow caused by vegetative dissemination of cultivated grapevine plants was not sufficient to genetically homogenise the pools of cultivars grown in different regions. The Neighbour Joining cluster analysis showed a clear separation according to geographical origins for the cultivated grapevines gene pools and revealed a high dissimilarity between cultivated and wild grapevine. However, three cultivars (Plant d’Ouchtata 1, Plant de Tabarka 3 and Plant d’Ouchtata 3) are very close to wild accessions and may result from a hybridisation between cultivated and wild accessions. The high level of differentiation between cultivated and wild accessions indicates that the cultivated accessions do not derive directly from local wild populations but could mostly correspond to imported materials introduced from others regions during historical times or derived from crossing between them.     

Population genetic structure of in situ wild Sorghum bicolor in its Ethiopian center of origin based on SSR markers

In situ population studies of wild relatives of crops are crucial for the conservation of plant genetic resources, especially in regions with high genetic diversity and a risk of local extinction. Ethiopia is the center of origin for sorghum, yet little is known about the genetic structure of extant wild populations. Using 9 Simple Sequence Repeat loci, we characterized 19 wild populations from five regions, 8 local cultivar populations from three regions, and 10 wild sorghum accessions from several African countries. To our knowledge, this is the most comprehensive study to date of in situ wild sorghum populations in Africa. Genetic diversity corrected for sample size was significantly greater in the wild populations in situ than in local cultivars or the accessions. Approximately 41 % of the genetic variation in the wild plants was partitioned among populations, indicating a high degree of differentiation and potential value for germplasm conservation, and the average number of migrants (N_m) per generation was 0.43. Cluster analyses showed that some wild populations were grouped by geographic region, whereas others were not, presumably due to long-distance seed movement. Four wild populations from disjunct regions formed a unique cluster with an Ethiopian accession of subsp. drummondii and probably represent a weedy race. STRUCTURE and other analyses detected evidence for crop-wild hybridization, consistent with previous molecular marker studies in Kenya, Mali, and Cameroon. In summary, in situ wild sorghum populations in Ethiopia harbor substantial genetic diversity and differentiation, despite their close proximity to conspecific cultivars in this crop/wild/weedy complex.     

Ecogeographical distribution of wild, weedy and cultivated Sorghum bicolor (L.) Moench in Kenya: implications for conservation and crop-to-wild gene flow

The potential gene flow between a crop and its wild relatives is largely determined by the overlaps in their ecological and geographical distributions. Ecogeographical databases are therefore indispensable tools for the sustainable management of genetic resources. In order to expand our knowledge of Sorghum bicolor distribution in Kenya, we conducted in situ collections of wild, weedy and cultivated sorghum. Qualitative and quantitative morphological traits were measured for each sampled wild sorghum plant. Farmers’ knowledge relating to the management of sorghum varieties and autecology of wild sorghum was also obtained. Cluster analysis supports the existence of several wild sorghum morphotypes that might correspond to at least three of the five ecotypes recognized in Africa. Intermediate forms between wild and cultivated sorghum belonging to the S. bicolor ssp. drummondii are frequently found in predominantly sorghum growing areas. Crop-wild gene flow in sorghum is likely to occur in many agroecosystems of Kenya.     

Genetic diversity in wild and cultivated black raspberry (Rubus occidentalis L.) evaluated by simple sequence repeat markers

Breeding progress in black raspberry (Rubus occidentalis L.) has been limited by a lack of genetic diversity in elite germplasm. Black raspberry cultivars have been noted for showing very few phenotypic differences and seedlings from crosses between cultivars for a lack of segregation for important traits. Despite these challenges, little molecular work has been done to explore genetic diversity and relationships in wild and cultivated black raspberry germplasm. Microsatellite, or simple sequence repeat (SSR), markers are highly polymorphic codominant markers useful for studying genetic diversity, population genetics, genetic fingerprinting and other applications. We examined genetic diversity in 148 wild and cultivated black raspberry accessions using 21 polymorphic SSR markers. Black raspberry cultivars clustered tightly and showed higher than expected heterozygosity while that of wild accessions was low. Relationships between wild black raspberry accessions were poorly resolved and regional clusters were mostly absent from our analysis. Our results indicated that wild black raspberry germplasm is a relatively untapped resource available for future breeding.     

Antioxidant characterization of some Sicilian edible wild greens

Epidemiological studies have demonstrated that many antioxidants and the total antioxidant capacity (TAC) of the diet may protect against cancers and cardiovascular disease. Common fruits and vegetables are good sources of antioxidants, although in some Mediterranean areas traditional wild greens are responsible for a significant percentage of total dietary antioxidant intake. In the European Prospective Investigation into Cancer and Nutrition cohort of Ragusa (Sicily), a high number of subjects were found to frequently eat wild greens, including Sinapis incana and Sinapis nigra, Diplotaxis erucoides, Cichorium intybus, Asparagus acutifolius, and Borrago officinalis. On the basis of these observations, detailed characterization of single antioxidant components (i.e., polyphenols, carotenoids, chlorophylls, and ascorbic acid) and the TAC of these edible wild traditional plants was performed. The wild plants examined were found to be very rich in antioxidants, such as flavonoids and carotenoids, with high TAC values, suggesting that the importance of these vegetables, not only in the traditional but even in the contemporary diet, needs to be emphasized.     

Plant genetic resources of Lemnos (Greece), an isolated island in the Northern Aegean Sea, with emphasis on landraces

Lemnos Island is located in the Northern Aegean Sea and presents a rich biodiversity in plant genetic resources including wild species and crop landraces. Landraces have been cultivated under traditional agricultural systems and many have survived genetic erosion. They are mainly conserved in home gardens or in small fields, usually by elderly people, and are limited to local consumption. Two collecting expeditions were organized in Lemnos, a minor one in 2009 and a larger one in 2010. The results of the two current expeditions are discussed and compared with previous expeditions in the island. An overview is given on the crops cultivated, the landraces collected and those considered as lost, alongside with some information about their traditional uses. Finally, the importance of landraces in modern agriculture and preservation policies in Lemnos are discussed.     

ISSR Variation in Olive-tree Cultivars from Morocco and other Western Countries of the Mediterranean Basin

In olive tree (Olea europaea L.), 12 varieties (or cultivars) representing the main domesticated material used in Morocco and 19 olive cultivars used extensively in five countries of the western Mediterranean Basin, were analysed using inter-simple sequences repeat (ISSR) markers which had never been used previously for extensive discrimination of cultivars. Four selected primers produced a total of 26 polymorphic reproducible amplification fragments. Combinations of these ISSR markers allowed to identify 25 of the 31 cultivars. Two additional combinations were distinguished, each corresponding to three Moroccan cultivars or local varieties. Evidence of a multiclonal composition in the widely cultivated variety ‘Picholine marocaine’ was obtained by the identification of three genotypes within the four morphologically distinct clones analysed in the variety. In the UPGMA phenogram based on the proportion of shared ISSR fragments, five groups of cultivars were distinguished at the 40% critical value of similarity. Four of the groups contained varieties from various geographic origins, as the consequence of successive human migrations which favoured olive dispersion throughout the Mediterranean Basin. However, the fifth group gathered together 9 of the 12 Moroccan cultivars and very few cultivated clones from Greece and Spain. The results suggest that most of the Moroccan cultivars are closely related and likely originated from local domestication.     

Maintenance of Squash (<Emphasis Type="Italic">Cucurbita</Emphasis> spp.) Landrace Diversity by Farmers' Activities in Mexico

Squash (Cucurbita spp.) is a common component in traditional cropping systems in Mexico, mainly in the agroecosystem known as the “milpa”, in which squash is cultivated in association with maize (Zea mays), the main crop. Using a questionnaire, 80 farmers were interviewed about crop production and selection practices in order to understand how these factors affect genetic diversity of local squash populations. We found that the most of the farmers who cultivate squash were elderly 59.8 ± 14.5 (mean ± SD; n = 78) years old. Squash varieties in the area were exclusively locally adapted landraces, and had not been replaced by modern squash cultivars. Two cultivated squash species, C. argyrosperma ssp. argyrosperma and C. moschata, had been grown intercropped with maize by 97.5% of the interviewed farmers, but only 50.0% were still producing squash at the time of the study. Farmers recognize typical characteristics of particular varieties within each of the local cultivated squash species, and selection is directed to maintain their identity. Nearly two thirds of the farmers (62.0%) had exchanged seeds of squash for planting, a practice that serves to increase genetic variability in the populations. All of the interviewed farmers were conscious of the possible hybridization between the wild gourd (C. argyrosperma ssp. sororia) and their cultivated squash. Despite various natural and human managed factors identified as contributing to enhancement of genetic diversity in these populations, results of the study show that genetic erosion of Cucurbita is likely in the region in the near future.