Breeding for increased wheat yields in a moisture-limited Mediterranean environment Abstract This literature review paper will focus

on covering major topics in increasing wheat yields, as part of long-term strategic solutions. NPK-style approach, organic farming and other hypotheses will be explained, as well as direct strategic methods for yield improvement will be reviewed: fertilization, selection, countering the effects of ultraviolet stress, managing factors like pests/parasites and genetic engineering for wheat culture improvement. The material presented is intended to fit into a coherent picture, briefly but extensively introducing an unfamiliar user into the subject matter, while certain topics regarding a moisture-limited Mediterranean environment are being discussed in parallel with generic ones, regarding all environments.

Introduction Wheat is one of the most important crops in the world, consisting mainly of two types hexaploid bread wheat (Triticum aestivum) and tetraploid hard (durum) wheat (T. durum). A typical Mediterranean diet includes a lot of meals that are made from both wheat types, so high yield production and further consumption of wheat is of high importance in related countries. Today, wheat ranks rst in world grain production and accounts for more than 20% of total human food calories. Wheat is extensively grown on 17% of all crop areas, in the temperate, Mediterranean-type and subtropical parts of both hemispheres, from 67N in Norway, Finland and Russia to 45S in Argentina. It is the staple food for 40% of the worlds population, mainly in Europe, North America and the western and northern parts of Asia (Peng et al., 2011). Improving yields of wheat is of high importance not only economically, but also to counter the effects of the recent century's explosion in human population, since our agricultural economies depend on domesticated crops, capable of surviving under cultivation in human agricultural niches (Peng et al., 2010). Nutrients and fertilization yield improvement strategies Nutrients are important for any crop growth. According to NPK hypothesis in agriculture, these three major elements Sodium, Phosphorous and Potassium are most essential for any culture rapid biomass growth. In case of wheat, most studies are centred around proper timing of applying sufficient amounts of N (Guoping et al., 2001; Nannipieri et al., 1999), its translocation in wheat plants under draught conditions (Xu et al., 2006) as well as ammonia exchange of between agricultural cropland and atmosphere, as a major pollutant (Schjoerring & Mattsson,

2001). Unfortunately, Raun et al. (2011) concluded that Nitrogen response and yield potential of crops (amount of yielded seeds per biomass) are quite unrelated as factors, so they both have to be included into the model estimating the amount of N to be applied, as well as timing of this application. Moreover, increasing evidence is being collected that application of N and other techniques as part of industrial growth of crops may not be sustainable in long term, in measures of yields (reviewed later). Regarding Mediterranean environment, a study by Xu et al. (2006) has concluded that nitrogen, especially excessively applied externally, gets translocated into grains of winter wheat under dry conditions (typical for mentioned environment). While in certain food industry applications this may or may not play a role, the flour product resulting from such grains may be inferior compared to normal one. A study by Nannipiery et al. (1999) on the other hand, concludes that the uptake of N by crops, as well as its recovery and retention is highly dependent on the various types of soils, generally present in Mediterranean environments. Moreover, in certain types of soil, like Foggia silty clay, the turnover of native soil N, being replaced by fertilizer N is less likely to happen, compared to Rielti soils. This introduces important strategic decisions to be made, when applying the fertilizer, as well as choosing its quality level and amounts. For example, if the synthetic fertilizer, contaminated by traces of chemical synthesis by-products gets retained in the soil, the latter will get contaminated and may become useless in the future. On the other hand, if N turnover rate is high, the certain soil type can be used to get a short-term strategic gain of a lower quality product for some specialized markets. Furthermore, addition of organic substances to soils has been shown beneficial to wheat yields (Barzegar et al., 2002), so such high turnover soils can be restored quickly, as compared to the more susceptible ones to contaminants.

Selection and shuttle breeding, the international wheat improvement strategy Selection and shuttle breeding is one of the major techniques applied to wheat, as part of the international wheat improvement strategy, developed by Mexican Centro Internacional de Mejoramiento de Maz y Trigo (CIMMYT), also known as the International Maize and Wheat Improvement Center. The first experiments in selection performed by CIMMYT were done in two locations in Mexico, with conditions probably quite similar to Mediterranean, except that the locations were in the Southern hemisphere. Selected wheats were shuttled between both locations quite often, to develop stem rust resistance first, but produced a completely new kind of wheat insensitive to day length (photoperiod), semidwarf, high yielding and disease-resistant (because crops were exposed to different disease spectra, soil types and environmental constraints in contrasting locations). This breakthrough has became known as the Green revolution, done by Borlaug and his colleagues. It is predicted by selection theory, that breeding a target population produces positive selection results only when yields of genotypes across various environments are positively correlated then selection in one environment may also improve yield in the correlated environment (Table 1). Factors and ranges Wide range of distinct environments Broad genetic background Low heritability due to significant genotype-byenvironment interaction and unreliable ranking of genotypes across environments. Narrow genetic background Maximizing variance among environments and significant means between testing environments.

Uniform environments.

Maximizing genetic variation and signicant means between testing genotypes.

Useless?

Table 1: Selection theory predictions according to ranges of environments and genotypes. (Ortiz, et al., 2007) Further selection strategy applied by CIMMYT in their research was to segregate selected cultures, and perform selection within specific regions, to enhance special features of existing culture strains. The new international wheat improvement strategy lies in applying the mentioned previous experience in shuttle breeding to an international wheat improvement effort, by shuttling wheat cultures across several continents worldwide. The program includes: introduction of new and novel sources of variation through wild species and, potentially, the use of transgenes for intractable traits; international shuttle breeding; enhanced breeding efciency through marker aided methods and more targeted use of physiology, genetics, statistics, and simulation to improve the design of crosses and early generation selection; an increased focus on the needs of end-users by emphasizing regional efforts in participatory research and client-oriented plant breeding. (Ortiz et al., 2007) When applied to Mediterranean territories, shuttle breeding can be done locally, facilitating germplasm exchange between quite adjacent regions to develop tolerance to certain pests or weeds, improve yields across various rainfall patterns, as well as internationally, as part of the new international wheat improvement strategy participation.

On the other hand, there is certain criticism of selection breeding in scientific literature, for example Calderini & Slafer conclude that genetic improvement of wheat yield potential has brought about an important decrease in absolute yield stability (Calderini & Slafer, 1999). According to these authors, while breeding for higher yields, the wheat culture automatically introduces capacity to respond to better environments. Probably, this quoted conclusion about absolute yield stability decrease may also get extended on genetic engineering of wheat cultures as well, since selection breeding is agigantic evolutionary experiment of adaptation and speciation, generating incipient species (Peng et al., 2011). Genetic engineering and analysis of wheat Genetic analysis of wheat genome has provided a lot of insight into processes of wheat domestication and selection. The domestication has been found to be restricted to a certain limited number of chromosome regions, termed domestication syndrome factors. The genome of diploid wild wheat (T. urartu or Ae. Tauschii) has been almost entirely sequenced, and research is now continued into mapping polyploidy plant species, by the International Wheat Genome Sequencing Consortium (IWGSC). Genetic engineering is sceptically accepted by international communities, especially when it is related to crop species or animals. By means of introducing alien genome sequences into existing genotype, special tolerant or resistant strains of wheat can be created ones that can theoretically survive lower winter temperatures, harsher dry conditions and compete more aggressively against weeds and pests. Unfortunately, not much of the existing developments can be applied to specifically Mediterranean environments. It is also important to note, that producing genetically modified wheat strains does not readily produce higher yields, since the

mechanisms for tolerance or competitiveness created by these manipulations can somehow influence other crop properties, in a manner that is not immediately obvious. It is also very important to preserve isolation of the natural course of wheat evolution and synthetic one, to as much extent as it is possible. Mostly, use of transgenes in wheat breeding is still at its inception (Ortiz et al., 2007)..

Countering other factors influencing yield: UV-B stress, pests and parasites Ultraviolet radiation generally badly affects growth of plants, because of oxidative damage stress. To sustain biomass growth rate, the existing antioxidant protective mechanisms of plants have to be enabled and/or facilitated, as well as other amelliorants or stress alleviants can be applied in parallel. The planetary ozone layer is depleted and estimated to lose 3% annually. Despite this questionable estimations, the UV radiation is still nevertheless present, and it is only rational, that facilitating plant defense mechanisms against existing normal or excess UV radiation could theoretically produce higher yields. The wheat's protection mechanism is rather complex and depends on a set of external macro elements and biochemistry inside the plant itself - generally activity of peroxidase (POD) and superoxide dismutase (SOD) and various catalase (CAT) enzymes activity, which influence the amount of superoxide radical (O2), and malondialdehyde (MDA) content regulation within the plant. Moreover, synthesis of compounds like anthocyanins, plavonoids and phenolics is essential for plant protection. It has been shown by Yao et al. (2010) that optimal supply of selenium (Se) microelement provides a short-strategy positive effect on growth and antioxidant traits of wheat (Triticum aestivum L. cv Han No.7086) seedlings exposed to enhanced UV stress (Figure 1).

Figure 1: Effect of Se suppply on growth of wheat seedlings exposed to UV-B radiation. (Yao et al., 2010)

Moreover, it has to be noted, that probably many other nutrients present in the soil, apart from the main NPK supplies, have a profound effect on plant's ability to protect itself from ultraviolet, by synthesizing and enabling appropriate enzymes and antioxidants, like phenolic compounds, in adequate quantities. More thorough studies of the protection mechanisms and their dependence/response to presence of various nutrients in the soil have to be done, while, currently the known optimal 1.0 mg kg-1 external supply of selenium can be applied as a shortterm strategy to counter the effects of ozone depletion. Pest eradication in industrial approach is usually done by pesticides and herbicides or by manual torch burning in special organic type farming (Pollan, 2007). In the former case, the yield of wheat that is partially sensitive to the chemical employed for pest inhibition can be affected, that is why extensive research is being done into selection and/or genetic engineering of

tolerant species (covered in chapters above). In Mediterranean cereal crop systems, one of the most significant pests types are the cereal aphids and parasites related to them (Lumbierres et al., 2007). Table 2 lists a comparison of various aphid and parasitoid species present in cereals, and it can be easily seen that wheat is one of the most vulnerable ones. Crop Wheat Aphids Rp, Sa, Md, Sm Parasitoids Ae, Am, Ar, Lt, Pv, Dr, Au, Aa, Aph Barley Rp, Sa, Md, Rm, Sg, Dn Oat Rye-grass Rp, Sa, Md Rp Ae, Am, Ar, Lt, Pv, Dr, Pg, Aph Ae, Am, Ar, Lt, Pv, Au, Aph Ae, Am, Ar

Dn Diuraphis noxia, Md Metopolophium dirhodum, Rm Rhopalosiphum maidis, Rp Rhopalosiphum padi, Sa Sitobion avenae, Sg Schizaphis graminum, Sm Sipha maydis, Aa Adialytus ambiguus, Ae Aphidius ervi, Am Aphidius matricariae, Ar Aphidius rhopalosiphi, Au Aphidius uzbekistanicus, Dr Diaretiella rapae, Lt Lysiphlebus testaceipes, Pg Praon gallicum, Pv Praon volucre, Aph Aphelinus sp.

Table 2: Aphid and parasitoid associated species for various cereal crops. (Lumbierres et al., 2007) In Catalonia (NE Iberian peninsula), aphids can cause direct damage to winter cereals (Lumbierres et al., 2007), as well as spring season. The main reason for this damage is theorized

to be the relatively short period between flowering and seed ripening and high temperatures during that period. The aphid-parasitoid interactions are rather complicated, as well as many other species prey for aphids, and this regulates their populations. In many cases the existing aphid populations are reservoirs for their parasitoids, and a strategy of aphid overwintering can be applied, allowing parasitoid-aphid interaction early in spring, which creates good natural control of aphids in winter. Unfortunately, not all aphid species are particularly vulnerable, or mechanisms of controlling them are not transparent. Natural predators, on the other hand, generally control the population outbreaks of aphids in spring seasons, so, theoretically some incentives may be possibly created for these birds to attend crop fields during pre-spring seasons.

Grass farming approach and related claims Grass farming is an approach to agriculture that is different from mainstream industrial or industrial organic. According to Michael Pollan (2007), a well known journalist and nutritionist, this approach is integrative to all aspects of crop and farming improvements, and provides better average sustained yields than conventional industrial methods, but, unfortunately little or no research confirming these claims has been made so far. The general idea of the approach is that big fields of domestic cultures are not natural and essentially are vulnerable to all kinds of attacks from pests, insects and other factors reducing yield. Moreover, because the soil employed for this kind of production gradually deteriorates (in terms of spectra of available chemicals and compounds, especially if fertilized purely or almost entirely according to NPK hypothesis) , the crops are unable to employ their mechanisms of self-protection to their full extent. The approach of grass farming states that large areas of land should be devoted to growing/farming ecosystems of grass and all kinds of living organisms ranging from bacteria and worms up to birds, cows and

other big animals. These ecosystems, on the other hand, will provide enough resources to fertilize and protect the crops, as well as restore the soil for future use, in rotating cycles. The industrial puritan ideas that nearly all pests can be eradicated from the fields, increasing yields are highly criticized, but, on the other hand, they are replaced by an idealistic claim that the nature itself, accompanied by smart farming can create entire ecosystems that take care of their own, and produce a sustainable way to support human needs. The author will abstain from any speculations for or against any of the farming ideologies, but entirely failing to mention other existing approaches, apart from mainstream, would seem to make the current literature review inferior, in terms of being complete.

Conclusions A brief review has been produced, covering a broad area of topics in wheat yield improvements. The approach to farming and selection of crops seems to be highly dependent on certain previous assumptions taken by the individual farmer or researcher as for granted, and then pursuing one's goals from these standpoints. Unfortunately, given the complexity of current problem, especially worldwide, one can not apply academic rigor to such research up to its full extent, and any results would probably involve mystery and guessing, as well as controversy surrounding them (even quite rigorous genome analysis). So, it is essential to cover the whole spectra of approaches and methodologies in most major environments geographic and academic, starting from literature reviews like this one.

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Calderini, D. & Slafer, G., 1999. Has yield stability changed with genetic improvement of wheat yield? Euphytica, 107(1), pp.5159. Peng, J., Sun, D. & Nevo, E., 2011. Domestication evolution, genetics and genomics in wheat. Molecular Breeding, 28(3), pp.281301. Yao, X., Chu, J. & Ba, C., 2010. Antioxidant Responses of Wheat Seedlings to Exogenous Selenium Supply Under Enhanced Ultraviolet-B. Biological Trace Element Research, 136(1), pp.96105. Pollan, M. (2007). Omnivore's Dilemma, Part II. Pastoral Grass. New York: Penguin Press.