Nutrient utilization efficiency (NUE): The case of nutrient use efficient genotypes

World population is expected to increase from 6.0 billion in 1999 to 8.5 billion by 2025. Such an increase in population will intensify pressure on the world’s natural resource base (land, water, and air) to achieve higher food production. Increased food production could be achieved by expanding the land area under crops and by increasing yields per unit area through intensive farming. Chemical fertilizers are one of the expensive inputs used by farmers to achieve desired crop yields [40]. However, during the last years, the prices of fertilizers have been considerably increased. Furthermore, soil degradation and pollution, as well as underground water pollution, are serious consequences provoked by the exaggerate use of fertilizers during last decades. These two aspects are responsible for the global concern to reduce the use of fertilizers. The best way to do that is by selecting and growing nutrient use efficient genotypes. According to Khoshgoftarmanesh (2009) [41], cultivation and breeding of micronutrient-efficient genotypes in combination with proper agronomic management practices appear as the most sustainable and cost-effective solution for alleviating food-chain micronutrient deficiency.

Nutrient use efficient genotypes are those having the ability to produce high yields under conditions of limited nutrient availability. According to Chapin and Van Cleve (1991) [11] and Gourley et al. (1994) [42], as nutrient utilization efficiency (NUE) is defined the amount of biomass produced per unit of nutrient absorbed. Nutrient efficiency ratio (NER) was suggested by Gerloff and Gabelman (1983) [43] to differentiate genotypes into efficient and inefficient nutrient utilizers, i. e. NER=(Units of Yields, kgs)/(Unit of elements in tissue, kg), while Agronomic efficiency (AE) is expressed as the additional amount of economic yield per unit nutrient applied, i. e. AE=(Yield F, kg-Yield C, kg)/(quantity of nutrient applied, kg), where F applies for plants receiving fertilizer and C for plants receiving no fertilizer.

Many researchers found significant differences concerning nutrient utilization efficiency among genotypes (cultivars) of the same plant species [1,12,13,40,44-46] Biomass (shoot and root dry matter production) was used as an indicator in order to assess Zn efficient Chinese maize genotypes, grown for 30 days in a greenhouse pot experiment under Zn limiting conditions [1]. NUE is based on: a) uptake efficiency, b) incorporation efficiency and c) utilization efficiency [40]. The uptake efficiency is the ability of a genotype to absorb nutrients from the soil; however, the great ability to absorb nutrients does not necessarily mean that this genotype is nutrient use efficient. According to Jiang and Ireland (2005) [45], and Jiang (2006) [46], Mn efficient wheat cultivars own this ability to a better internal utilization of Mn, rather than to a higher plant Mn accumulation. We also found in our experiments that, despite the fact that the olive cultivar ‘Kothreiki’ absorbed and accumulated significantly greater quantity of Mn and Fe in three soil types, compared to ‘Koroneiki’, the second one was more Mn and Fe-efficient due to its better internal utilization efficiency of Mn and Fe (greater transport of these micronutrients from root to shoots) [12] (Tables 1 and 2). Aziz et al. (2011a) [47] refer that under P deficiency conditions, P content of young leaves in Brassica cultivars increased by two folds, indicating remobilization of this nutrient from older leaves and shoot. However, differences in P remobilization among Brassica cultivars could not explain the differences in P utilization. Phosphorus efficient wheat genotypes with greater root biomass, higher P uptake potential in shoots and absorption rate of P were generally more tolerant to P deficiency in the growth medium [6]. According to Yang et al. (2011) [48], on average, the K efficient cotton cultivars produced 59% more potential economic yield (dry weight of all reproductive organs) under field conditions even with available soil K at obviously deficient level (60 mg/kg).

The possible causes for the differential nutrient utilization efficiency among genotypes and/or species may be one, or combination of more than one, of the following: a) genetic reasons (genotypic ability to absorb and utilize efficiently, or inefficiently, soil nutrients), b) mycorrhiza colonization of the root system, c) differential root exudation of organic compounds favorizing nutrient uptake, d) different properties of rhizosphere, e) other reasons. According to Cakmak (2002) [49], integration of plant nutrition research with plant genetics and molecular biology is indispensable in developing plant genotypes with high genetic ability to adapt to nutrient deficient and toxic soil conditions and to allocate more micronutrients into edible plant products. According to Aziz et al. (2011b) [50], Brassica cultivars with high biomass and high P contents, such as ‘Rainbow’ and ‘Poorbi Raya’, at low available P conditions would be used in further screening experiments to improve P efficiency in Brassica. More specifically, a number of genes have been isolated and cloned, which are involved in root exudation of nutrient-mobilizing organic compounds [51,52]. Successful attempts have been made in the past 5 years to develop transgenic plants that produce and release large amounts of organic acids, which are considered to be key compounds involved in the adaptive mechanisms used by plants to tolerate P-deficient soil conditions [53-55]. However, differential root exudation ability in nature exists among different plant species. According to Maruyama et al. (2005) [56], who made a comparison of iron availability in leaves of barley and rice, the difference in the Fe acquisition ability between these two species was affected by the differential mugineic acid secretion. Chatzistathis et al. (2009) [12] refer that, maybe, a similar mechanism was responsible for the differential micronutrient uptake and accumulation between the Greek olive cultivars ‘Koroneiki’ and ‘Kothreiki’. According to the same authors, differential reduction of Fe3+ to Fe2+, or acidification capacity of root apoplast (which associates with the increase of Fe3+-chelate reductase and H-ATPase activities) among three Greek olive cultivars should not be excluded from possible causes for the significant differences observed concerning Fe uptake [14]. Mycorrhiza root colonization may be another responsible factor for the differential micronutrient utilization efficiency among genotypes. According to Citernesi et al. (1998) [57], arbuscular mycorrhiza fungi (AMF) influenced root morphology of Italian olive cultivars, thus nutrient uptake and accumulation, as well as plant growth. In our study with olive cultivars ‘Koroneiki’, ‘Kothreiki’ and ‘Chondrolia Chalkidikis’, we found significant differences concerning root colonization by AMF (that varied from 45% to 73%), together with great differences in uptake and utilization efficiency of Mn, Fe and Zn among them (particularly, 1.5 to 10.5 times greater amount of Mn, Fe and Zn accumulated by ‘Kothreiki’, compared to the other two cultivars, but the differences in plant growth parameters between the three cultivars were not impressive; this is why the micronutrient utilization efficiency by ‘Kothreiki’ was significantly lower, compared to that of the other two ones). Finally, the different properties of rhizosphere among genotypes may be another important factor influencing nutrient uptake and utilization efficiency, and of course biomass production. According to Rengel (2001) [58], who made a review on genotypic differences in micronutrient use efficiency of many crops, micronutrient-efficient genotypes were capable of increasing soil available micronutrient pools through changing the chemical and microbiological properties of the rhizosphere, as well as by growing thinner and longer roots and by having more efficient uptake and transport mechanisms.

Soil

Cultivar

Micronutrient

Root

Stem

Leaves

Marl

Mn

Kor

50.2b

38.0a

11.8a

Koth

74.1a

12.8b

13.1a

Gneiss

schist

Kor

56.5b

34.2a

9.3a

Koth

81.3a

10.8b

7.9a

Peridotite

Kor

44.0b

44.0a

12.0a

Koth

76.0a

12.9b

11.1a

Marl

Fe

Kor

93.7a

3.9a

2.4a

Koth

98.0a

0.9b

1.1b

Gneiss

schist

Kor

94.0a

3.7a

2.3a

Koth

98.8a

0.6b

0.6b

Peridotite

Kor

90.8a

7.1a

2.1a

Koth

98.3a

0.8b

0.9b

Marl

Zn

Kor

49.3b

29.6a

21.1a

Koth

64.4a

15.6b

20.0a

Gneiss

schist

Kor

59.1b

26.7a

14.2a

Koth

73.7a

14.3b

12.0a

Peridotite

Kor

37.3b

33.9a

28.8a

Koth

65.3a

18.0b

16.7b

The different letters in the same column symbolize statistically significant differences between the two olive cultivars in each of the three soils, for P<0.05 (n=6) (SPSS; t-test).

Table 1. Distribution (%) of the total per plant quantity of Mn, Fe and Zn in the three vegetative tissues (root, stem and leaves) of the olive cultivars ‘Koroneiki’ and ‘Kothreiki’, when each one was grown in three soils (from parent material Marl, Gneiss schist. and Peridotite) with different physicochemical properties (Chatzistathis et al., 2009).

Soil

Cultivar

MnUE

FeUE

ZnUE

Marl

mg of the total plant d. w./pg of the total per plant quantity of micronutrient

Kor

31.85a

1.73a

77.53a

Koth

18.68b

0.65b

68.08a

Gneiss schist

Kor

39.87a

1.84a

51.04a

Koth

17.94b

0.44b

49.15a

Peridotite

Kor

23.33a

1.19a

61.75a

Koth

18.00a

0.58b

72.88a

The different letters in the same column symbolize statistically significant differences between the two cultivars in each of the three soils, for P<0.05 (n=6) (SPSS; t-test).

Table 2. Nutrient utilization efficiency (mg of the total plant d. w. /^g of the total per plant quantity of micronutrient or mg of the total per plant quantity of macronutrient) of the olive cultivars ‘Koroneiki’ and ‘Kothreiki’, when each of them was grown in three soils (from parent material Marl, Gneiss schist. and Peridotite) with different physicochemical properties (Chatzistathis et al., 2009).