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Mineral Nutrition of Plants

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DOI: 10.1007/978-81-322-2286-6_20

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Chapter Title Mineral Nutrition of Plants

Copyright Year
Copyright Holder
Corresponding Author
Abstract
Keywords (separated by “ - ”)
2015
Springer India
Family Name Pandey
Particle
Given Name Renu
Suffix
Division Mineral Nutrition Laboratory,
Division of Plant Physiology
Organization/University Indian Agricultural Research
Institute
City New Delhi
Postcode 110012
Country India
Email renu.pandey.iari@gmail.com
In this chapter, a brief overview of the history of plant mineral
nutrition is provided. Soil serves as the source of nutrient elements,
and so the availability of nutrients is governed by soil properties.
The term ‘essential mineral element’ has been defined, and these
elements are grouped according to their biochemical behaviour
and physiological functions. In addition to these elements, another
group of elements called beneficial elements has been discussed.
The pathway of movement of elements (as ions) through roots and
different mechanisms for absorption and transport of nutrients has
been outlined in this chapter. Besides the inorganic nutrient elements,
the use of biofertilisers in agriculture has been discussed. These
biofertilisers associate with the plants either symbiotically or non-
symbiotically, to help them enhance absorption of nutrient elements
from soil and improve growth.
Plants - Mineral nutrition - Macro- and microelements - Essential
mineral element - Transport and absorption of nutrients - Biofertilisers
 Mineral Nutrition of Plants 1
20
Renu Pandey 2

Abstract 3
In this chapter, a brief overview of the history of plant mineral nutrition is 4
provided. Soil serves as the source of nutrient elements, and so the avail- 5
ability of nutrients is governed by soil properties. The term ‘essential min- 6
eral element’ has been defined, and these elements are grouped according 7
to their biochemical behaviour and physiological functions. In addition to 8
these elements, another group of elements called beneficial elements has 9
been discussed. The pathway of movement of elements (as ions) through 10
roots and different mechanisms for absorption and transport of nutrients 11
has been outlined in this chapter. Besides the inorganic nutrient elements, 12
the use of biofertilisers in agriculture has been discussed. These biofertil- 13
isers associate with the plants either symbiotically or non-symbiotically, 14
to help them enhance absorption of nutrient elements from soil and 15
improve growth. 16

Keywords 17
Plants • Mineral nutrition • Macro- and microelements • Essential mineral 18
element • Transport and absorption of nutrients • Biofertilisers 19

20.1 Introduction 20

Inorganic minerals present in the Earth’s crust are [AU1]

21
used for nutrition by plants by extracting them 22
from soil or the aquatic environment. These min- 23
eral elements are formed by the complex interac- 24
tion involving weathering of rock minerals, decay 25
R. Pandey (*) of organic matter, animals and microbes. Among 26
Mineral Nutrition Laboratory, Division of Plant nutrient elements, nitrogen is exceptional as its 27
Physiology, Indian Agricultural Research Institute,
New Delhi 110012, India primary source is gaseous nitrogen of the atmo- 28
e-mail: renu.pandey.iari@gmail.com sphere and little occurs in minerals. Roots absorb 29

B. Bahadur et al. (eds.), Plant Biology and Biotechnology: Volume I: Plant Diversity,
Organization, Function and Improvement, DOI 10.1007/978-81-322-2286-6_20,
© Springer India 2015
 R. Pandey

30 mineral nutrients in the form of their salts dis- Table 20.1  Discovery of essentiality of micronutrients t1.1
for higher plants (Marschner 1995) t1.2
31 solved in soil water. The study of absorption of
32 inorganic mineral elements and their assimilation Element Year Discovered by t1.3

33 by plants is called mineral nutrition. Once the Iron 1860 J. Sachs t1.4

34 elements are absorbed by roots, they are translo- Manganese 1922 J. S. McHargue t1.5

35 cated to various parts of the plant where they are Boron 1923 K. Warington t1.6

36 involved in carrying out important biological Zinc 1926 A. L. Sommer and C. B. t1.7
Lipman t1.8
37 functions resulting in normal growth and
Copper 1931 C. B. Lipman and t1.9
38 development. G. Mackinney t1.10
39 In agriculture, the addition of mineral ele- Molybdenum 1938 D. I. Arnon and P. R. Stout t1.11
40 ments to soil to improve plant growth dates back Chlorine 1954 T. C. Broyer et al. t1.12
41 to more than 2000  years. About 150  years ago, Nickel 1987 P. H. Brown et al. t1.13
42 the function of mineral nutrients in plant growth
43 was a topic of scientific debate. However, it was
44 Justus von Liebig (1803–1873) who collected, were carried out in which particular elements 78
45 compiled and summarised the scattered informa- were omitted. The technique of growing plants in 79
46 tion pertaining to the importance of mineral ele- soilless culture media was termed as hydropon- 80
47 ments for plant growth. This established the ics. These techniques were used to characterise 81
48 mineral nutrition of plant as a scientific disci- the essentiality of individual mineral elements 82
49 pline. These achievements led to a rapid increase more precisely and led to a better understanding 83
50 in the use of mineral fertilisers in agriculture. By of their role in plant metabolism. Discovery of 84
51 the end of the nineteenth century, particularly in essentiality of micronutrients by various workers 85
52 Europe, large amount of potash, superphosphate using hydroponics is presented in Table 20.1. 86
53 (phosphorus) and, later, inorganic nitrogen was
54 used in agriculture and horticulture to improve
55 plant growth. It was concluded based solely on 20.2 Soil as Source of Nutrient 87
56 observation and speculation rather than by pre-
57 cise experimentation that mineral elements such Soil is a complex physical, chemical and biologi- 88
58 as nitrogen, sulphur, phosphorus, potassium, cal- cal substrate which acts as a matrix for various 89
59 cium, magnesium, silicon, sodium and iron are organic and inorganic nutrients. Soil contains all 90
60 essential for plant growth. the three phases, viz., solid, liquid and gaseous, 91
61 By the end of nineteenth century, a large num- which interact with mineral elements. The solid 92
62 ber of studies were undertaken to establish the phase provides a reservoir of both inorganic 93
63 ‘mineral element theory’. From the extensive nutrients (potassium, calcium, magnesium and 94
64 investigation carried out on mineral composition iron) as well as organic compounds which pro- 95
65 of different plant species growing on different vide nitrogen, phosphorus and sulphur among 96
66 types of soils, it was concluded that neither the other elements. The liquid phase or the soil solu- 97
67 presence nor the concentration of mineral ele- tion is very important from the view point of 98
68 ment in a plant is the criteria for essentiality. absorption of nutrients by roots because it con- 99
69 Plants have a limited capability to absorb selec- tains dissolved nutrient ions and facilitates the 100
70 tively those mineral nutrients that are essential movement of ions towards root surface. The air- 101
71 for their growth. They also take up mineral ele- spaces between soil particles which are occupied 102
72 ments that are not essential for growth and may with gases such as oxygen, carbon dioxide and 103
73 be even toxic. Therefore, it was evident that the nitrogen constitute the gaseous phase of soil. 104
74 mineral composition of plants growing in soils From the biological perspective, soil constitutes a 105
75 cannot be used as a criterion to judge the essenti- diverse ecosystem in which plant roots and 106
76 ality of a mineral element. Once this fact was estab- microorganisms compete strongly for mineral 107
77 lished, both water and sand culture experiments nutrients. In spite of this competition, roots and 108
 20  Mineral Nutrition of Plants

109 microorganisms can form associations for their (leaching loss). However, these adsorbed nutrients 154
110 mutual benefit (nitrogen-fixing bacteria, arbuscu- can be replaced by other cations in a process 155
111 lar mycorrhiza). known as cation exchange. The capacity of a soil 156
112 Based on the bioavailability of inorganic to adsorb ions and exchange it with other ions is 157
113 nutrient elements to the plant roots, the soil may termed as cation exchange capacity (CEC) and is 158
114 be considered as fertile or non-fertile. The term highly dependent on soil types. Soils with small 159
115 fertility refers to the inherent capacity of soil to particles like clays have a high ratio of surface 160
116 supply nutrients to plants in adequate amounts area to volume resulting in a higher CEC. A soil 161
117 and appropriate proportions. In non-fertile soils, with higher CEC generally has larger reserves of 162
118 in order to increase the availability of nutrients in mineral nutrients. On the other hand, mineral 163
119 a balanced proportion, inorganic elements in the anions such as nitrate (NO3−), chloride (Cl−) and 164
120 form of fertilisers are added externally. Most fer- phosphates (PO43−) do not get adsorbed because 165
121 tilisers are formulated to overcome the deficien- of repulsion due to the negative charge on the sur- 166
122 cies of mineral elements in soil. face of soil particles and thus remain dissolved in 167
the soil solution. These anions are highly prone 168
to leaching loss. 169
123 20.2.1 Soil Properties Affecting Another important chemical property affect- 170
124 Nutrient Availability ing nutrient availability is pH or the hydrogen ion 171
concentration of soil solution. The reaction of 172
125 The chemical (surface charge, pH) and physical soil solution, whether it is neutral, alkaline or 173
126 (soil texture and structure) properties of soil acid, has a distinct effect on the availability of 174
127 determines the availability of nutrients on the mineral elements to plant roots. The range of pH 175
128 root surface. The soil particles, both organic and for most crops lies approximately between 5.5 176
129 inorganic, are negatively charged. This negative and 6.5 at which the greatest average levels of all 177
130 charge is due to the fact that many inorganic soil essential plant nutrients become available 178
131 particles are crystal lattices which are arranged in (Fig. 20.1). This pH range also favours the root 179
132 a tetrahedral form of aluminium and silicon (Al3+ growth. Extreme fluctuations of soil pH on either 180
133 and Si4+). These tetrahedral arrangements are side can cause nutrient imbalances in plants 181
134 covalently bound to oxygen atoms forming alu- resulting in deficiency or toxicity symptoms. 182
135 minates and silicates. These tetrahedral struc- The physical property of soil includes differ- 183
136 tures undergo isomorphous substitution wherein ent sizes of soil particles. The relative proportion 184
137 another cation of lesser charge replaces the Al3+ of various soil particle types is referred to as soil 185
138 or Si4+ thus making them negatively charged par- texture. The arrangement of soil particles, that is, 186
139 ticles. Microbial decomposition of dead plants, sand, silt and clay in soil, is known as soil struc- 187
140 animals and microorganisms leads to the forma- ture. The soil particles are named as gravel, 188
141 tion of organic soil particles. The dissociation of coarse sand, fine sand, silt and clay based on two 189
142 hydrogen ions from carboxylic acid and phenolic systems as shown in Table 20.2. Practically, all 190
143 groups present in organic soil particles provides soils are mixtures of sand, silt and clay. Soils 191
144 negative surface charges. with 10–25 % clay and the rest about equal parts 192
145 Charge on the surface of soil particles has an of sand and silt are called loams. The soil texture 193
146 important role in plant nutrition. Mineral cations governs the fertility of soil. 194
147 such as ammonium (NH4+) and potassium (K+) Soil aeration is another important physical 195
148 adsorb to the negatively charged surface of the property affecting the availability of some of the 196
149 organic and inorganic soil particles. This adsorp- nutrient elements. Soils rich in clay and humus 197
150 tion is an important factor in governing soil fertil- can hold more water expelling air from the space 198
151 ity. The major advantage of adsorption is that the between the particles. Such soil lacks oxygen and 199
152 cations are not easily lost from the surface of soil is not ideal for plant growth. In these waterlogged 200
153 particles when the soil is drained with water (reduced) soils, the availability of reduced elements 201
 R. Pandey

Fig. 20.1 Relationship
between level of availabil-
ity of different elements
and soil pH (Source:
Goedert et al. 1997)

t2.1 Table 20.2  Classification of soil particles by diameter 1. A plant must be unable to complete its life 222
t2.2 (Salisbury and Ross 1986)
cycle in the absence of the mineral element. 223
t2.3 USDA system 2. The function of the element must not be 224
t2.4 Particle (mm) World system (mm)
replaceable by another mineral element. 225
t2.5 Coarse sand 2.0–0.2
3. The element must be directly involved in plant 226
t2.6 Fine sand 2.0–0.05 0.2–0.02
metabolism, for example, as a component of 227
t2.7 Silt 0.05–0.002 0.02–0.002
an essential plant constituent such as an 228
t2.8 Clay <0.002 <0.002
enzyme or it must be required for a distinct 229
metabolic step such as an enzymatic reaction. 230
The second criterion has some exceptions; it 231
202 such as Fe3+ increases. The microorganism pres- cannot be followed as such because there are 232
203 ent in waterlogged soil utilises Fe2+ as an electron some elements that can be replaced by others 233
204 acceptor and thus Fe is reduced. This reduced without causing any adverse effect on the plant. 234
205 form of Fe is absorbed in excess by some plants For example, the monovalent cation K+ can be 235
206 adapted to wetland such as paddy and suffer from replaced by Na+; both these ions play important 236
207 iron toxicity (bronzing). The best agricultural soil roles in osmoregulation. On the other hand, this 237
208 from the point of view of good plant growth and strict definition of essentiality excludes those 238
209 nutrient availability are sandy loams and clay mineral elements that compensate for toxic 239
210 loams. effects of other elements or replace mineral nutri- 240
ents involved in some specific function. Such ele- 241
ments are not essential but perform certain 242
211 20.3 C
 riteria for Essentiality important functions and are classified as benefi- 243
212 of Elements cial elements. By definition, beneficial elements 244
are those that stimulate growth but are not essen- 245
213 The nutrient elements essential for healthy tial or might be essential for certain plant species 246
214 growth of plants are called essential nutrients or under specific conditions. The optimum genetic 247
215 essential mineral elements. Till date, only 17 ele- potential of crop plants cannot be achieved if the 248
216 ments (nickel being the recently enlisted) are beneficial elements are excluded from the agri- 249
217 considered as essential. Arnon and Stout in the cultural production system. 250
218 year 1939 proposed the term ‘essential mineral Based on the essentiality criteria, mineral ele- 251
219 element’. They established that the following ments have specific and essential functions in 252
220 three criteria must be met for an element to be plant metabolism. Therefore, depending on the 253
221 considered essential. requirement of a nutrient element to produce 254
 20  Mineral Nutrition of Plants

255 optimum plant growth, the nutrient is referred to nutrients into metals (K, Ca, Mg, Fe, Mn, Zn, Cu, 270
256 as either macronutrient or micronutrient. The Mo, Ni) and non-metals (N, S, P, B, Cl). However, 271
257 macronutrients are required in larger quantities the most widely used classification is based on 272
258 and are present in plant tissues in amounts rang- the quantity of requirement of mineral element 273
259 ing between 0.2 and 4.0  % (on a dry weight rather than the physiochemical properties. 274
260 basis), while the concentration of micronutrients
261 in plant tissue ranges from 5 to 200 ppm or less
262 than 0.02 %. 20.3.1 Quantitative Requirements 275
263 The macronutrients are further divided based of Nutrient and Tissue 276
264 on their requirements into primary macronutri- Analysis 277
265 ents consisting of nitrogen (N), phosphorus (P)
266 and potassium (K) and secondary macronutrients The concentrations of essential elements required 278
267 including calcium (Ca), sulphur (S) and magne- by plants for maintaining optimum growth and 279
268 sium (Mg) (Table  20.3). Another classification preventing any deficiency symptoms are given in 280
269 based on physiochemical properties divides Table 20.4. Such values serve as useful guide to 281
plant physiologists and farmers because concen- 282

t3.1 Table 20.3  Essentiality of mineral elements for higher tration of elements in plant tissue is more reliable 283
t3.2 plants than in soil indicating whether the plant will grow 284

t3.3 Higher faster if more of a given nutrient is applied. 285

t3.4 Classification Element plants However, for determining the optimum growth of 286
t3.5 Macronutrient N, P, S, K, Mg, Ca + a plant, a farmer should take into consideration 287
t3.6 Micronutrient Fe, Mn, Zn, Cu, B, + the soil analysis of his field. Soil analysis is the 288
t3.7 Mo, Cl, Ni chemical determination of the nutrient content 289
t3.8 Beneficial elements Na, Si, Co, Al ± and their quantification in soil sample. The soil 290

t4.1 Table 20.4  The form of nutrient taken up plant and its average concentration in shoot dry matter required for adequate
t4.2 growth
t4.3 Concentration in
t4.4 dry tissue
t4.5 Element Abbreviation Form taken by plants (ppm) (%) Relative no. of atoms
t4.6 Molybdenum Mo MoO4− 0.1 0.00001 1
t4.7 Nickel Ni Ni2+ ~0.1 0.00001 1
t4.8 Copper Cu Cu+, Cu2+ 6 0.0006 100
t4.9 Zinc Zn Zn2+ 20 0.002 300
t4.10 Manganese Mn Mn2+ 50 0.005 1,000
t4.11 Iron Fe Fe3+, Fe2+ 100 0.01 2,000
t4.12 Boron B H3BO3 20 0.002 2,000
t4.13 Chlorine Cl Cl− 100 0.01 3,000
t4.14 Sulphur S SO42− 1,000 0.1 30,000
t4.15 Phosphorus P H2PO4−, HPO42− 2,000 0.2 60,000
t4.16 Magnesium Mg Mg2+ 2,000 0.2 80,000
t4.17 Calcium Ca Ca2+ 5,000 0.5 125,000
t4.18 Potassium K K+ 10,000 1.0 250,000
t4.19 Nitrogen N NO3−, NH4+ 15,000 1.5 1,000,000
t4.20 Oxygen O O2, H2O 450,000 45.0 30,000,000
t4.21 Carbon C CO2 450,000 45.0 35,000,000
t4.22 Hydrogen H H2O 60,000 6.0 60,000,000
t4.23 Source: Modified after Salisbury and Ross (1986)
t4.24 Between one of the two forms of element, bold-faced type is the preferred form by plants
 R. Pandey

Fig. 20.2  Generalised plot al Adequate Zone

gin

M Zon
of growth rate as a function

ar e
100 ar ne

gi
of the concentration of a M Zo

Yield (% of maximum)

na
90

l
nutrient element in plant 90% Maximum Yield
tissue

one

Tox
nt Z
50

ic
ficie

Zon
De
Critical Concentrations

e
Deficiency Toxicity
0
Nutrient concentration in tissue

291 analysis reflects overall levels of nutrients poten- that sustain yield and ensure the food quality of 326
292 tially available to plant roots. However, it fails to many crops. 327
293 evaluate the uptake conditions and amount of The distribution of mineral nutrients between 328
294 nutrients actually absorbed by the plants. This different types of cells within a given tissue (e.g. 329
295 additional information can be obtained by plant epidermis, guard cells, mesophyll cells of leaf) 330
296 tissue analysis. also provides important information about func- 331
297 Proper use of plant tissue analysis requires an tion of mineral nutrients. This is particularly true 332
298 understanding of the relationship between plant for the distribution of ions in different cellular 333
299 growth and the mineral content of plant tissue compartments. In the last decade, much progress 334
300 samples. Figure 20.2 shows an idealised plot of has been made in this respect by applying tech- 335
301 growth rate as a function of the concentration of niques, such as X-ray microanalysis, nuclear 336
302 any given nutrient element in the plant. When magnetic resonance (NMR), ion-selective micro- 337
303 the nutrient content in tissue sample is low, electrodes or fluorescent dyes, for studies on ion 338
304 growth is reduced called the deficient zone. In distribution in cytoplasm and the organelles con- 339
305 this zone, an increase in tissue mineral content tained within it (e.g. chloroplast) and the vacuole. 340
306 is directly related to an increase in growth or New insight into the functions of mineral nutri- 341
307 yield. As the nutrient content in tissue sample ents, for example, of calcium as a secondary mes- 342
308 increases further, a point is reached at which senger, is based on these studies of cellular 343
309 additional increases in tissue mineral content do compartmentation. David E.  Salt and his group 344
310 not appreciably affect growth or yield. This (2008) have proposed the concept of ‘ionome’. 345
311 region of the curve is called adequate zone. The They defined ‘ionome’ as the entire mineral 346
312 transition between deficiency and adequate zone nutrient and trace element composition of an 347
313 of the curve represents the critical concentration organism representing the inorganic component 348
314 of the nutrient in question, which may be defined of cellular and organismal systems. Hence, ion- 349
315 as the minimum tissue concentration of the ome includes both essential and non-essential 350
316 nutrient that is sufficient to give maximal growth elements. Ionomics is the study of the ionome, 351
317 or yield. The adequate zone also represents the involving quantitative and simultaneous mea- 352
318 luxury consumption of elements during which surement of the elemental composition of living 353
319 there is no increase in growth and the nutrient organisms and changes in this composition in 354
320 taken up in excess is stored in vacuoles. This response to physiological stimuli, developmental 355
321 occurs for a few elements like K, otherwise cues and genetic modifications. This study 356
322 increases in tissue concentration beyond the requires application of high-throughput elemen- 357
323 adequate zone results in toxicity limiting the tal analysis technologies such as inductively cou- 358
324 growth or yield. Plant analysis and soil analysis pled plasma-optical emission spectroscopy 359
325 data are useful in establishing fertiliser schedules (ICP-OES) or ICP-mass spectroscopy (ICP-MS) 360
 20  Mineral Nutrition of Plants

361 and their integration with both bioinformatics 20.4.1 Physiological Functions 405
362 and genetic tools. The information about functional of Macronutrients 406
363 state of an organism under different conditions
364 brought about by the genetic and developmental 20.4.1.1 Nitrogen 407
365 differences and by biotic and abiotic factors can The Earth’s atmosphere consists of about 80  % 408
366 be captured by ionomics. N, but the extremely stable form of atomic N 409
367 The progress towards a better understanding (dinitrogen, N2) is not available to plants. 410
368 of the function of mineral nutrients helps in com- However, the microorganisms both free-living 411
369 paring genotypes or mutants within a given plant and symbiotic can fix atmospheric N2 in ammo- 412
[AU2] species. The advanced techniques and concept in
370 niacal form (NH4+) which is then directly taken 413
371 mineral nutrition like ionome analysis provides a up by the plants or converted into nitrate (NO3−) 414
372 powerful approach to not only analyse the func- by nitrifying bacteria. The preferred form in 415
373 tions of genes and gene networks directly con- which N is taken up depends on soil conditions 416
374 trolling the ionome but also the extended gene and plant species. Plants adapted to low pH and 417
375 networks that control physiological and develop- waterlogged soil conditions tend to take up NH4+, 418
376 mental processes which indirectly affects the for example, paddy. In aerobic soils with higher 419
377 ionome. pH, NO3− is the predominant form preferred by 420
most of the plants. Also, organic N compounds 421
such as amino acids are found in soil, and there is 422
378 20.4 I mportance of Macro growing evidence that these can also form impor- 423
379 and Microelements tant N sources. 424

380 For easy understanding, both the macro- and 20.4.1.1.1  N Uptake and Distribution 425
381 micronutrient elements can be classified into four The uptake of nitrate or ammonium into plants 426
382 groups as given below (Malik and Srivastava from soil via the roots involves two basic catego- 427
383 1982): ries of transport system, high-affinity transport 428
384 Group 1: N and S – present in reduced form and system (HATS) and low-affinity transport system 429
385 are covalently bonded constituents of organic (LATS), identified by kinetic studies of N uptake. 430
386 matter The HATS operates under conditions of low 431
387 Group 2: P, B and Si – occurs as oxyanions such external nitrate concentration (Km <200 μM), and 432
388 as phosphate, borate or silicate LATS operates under higher external concentra- 433
389 Group 3: K, Na, Mg, Ca and Cl  – involved in tions even as high as 50 mM without saturation. 434
390 osmoregulation and ionic balance and have Based on molecular, physiological and biochemi- 435
391 specific functions of enzyme conformation cal studies, the transport systems were further 436
392 and catalysis (e.g. metalloprotein complexes) subdivided into inducible and constitutive depend- 437
393 Group 4: Fe, Cu, Mo and Zn – present as struc- ing on substrate induction process. The HATS is 438
394 tural chelates or metalloproteins; also involved comprised of inducible and constitutive HATS 439
395 in oxidation-reduction (redox) reactions (first (iHATS and cHATS) and is encoded by the mem- 440
396 three elements) bers of NRT2 gene family (Williams and Miller 441
397 The following paragraphs describe the occur- 2001). The constitutive system (cHATS) is avail- 442
398 rence and physiological functions of macro- and able when plant has been previously starved for 443
399 micronutrients in detail with recent developments nitrate, and the inducible system (iHATS) is stim- 444
400 in the field of mineral nutrition. A summary of ulated by supplying nitrate. Similarly, LATS 445
401 physiological functions and the deficiency symp- encoded by members of NRT1 gene family is also 446
402 toms of mineral elements in plants are presented comprised of both constitutive and inducible ele- 447
403 in Table 20.5 (modified after Malik and Srivastava ments, evidence of which was shown in 448
404 1982). Arabidopsis thaliana. First inducible LATS gene 449
 R. Pandey

t5.1 Table 20.5  Summary of physiological functions and the deficiency symptoms of mineral elements in plants
t5.2 Elements Primary function Specific deficiency symptoms
t5.3 Oxygen Final electron acceptor in aerobic
t5.4 respiration; element in carbohydrates,
t5.5 nucleic acids and many other organic
t5.6 compounds
t5.7 Carbon Building block of all organic compounds
t5.8 in the plants’ body
t5.9 Hydrogen Component of water and all organic
t5.10 compounds
t5.11 Nitrogen Constituent of amino acids, proteins, ‘V’ shape chlorosis starting from the tip in
t5.12 chlorophyll, nucleic acid and some lower leaves, plant becomes pale green, poor
t5.13 coenzymes growth
t5.14 Potassium Involved in enzyme activation, protein Chlorosis of older leaves later turns into
t5.15 metabolism, cell membranes, ionic necrotic lesions on leaf tip, rolling of leaves,
t5.16 balance, cell extension growth, opening shortening of internodes leading to stunted
t5.17 and closing of stomata, cell turgor growth
t5.18 Calcium Constituent of cell wall, middle lamella, Twisting and deformation of growing tips and
t5.19 enzyme cofactor, controls cell youngest leaves, later necrosis occurs at the leaf
t5.20 permeability and cell wall configuration, margin
t5.21 involved in cell signalling
t5.22 Phosphorus Involved in photosynthesis, constituent of Stunted growth, foliage turns dark green, high
t5.23 high-energy intermediaries like ATP, root/shoot ratio, delayed maturity of plants
t5.24 nucleic acids, phospholipids in membranes
t5.25 Magnesium Central element of chlorophyll molecule Interveinal chlorosis of older leaves, purple
t5.26 and an activator of several enzymes coloration with necrotic spots, leaves become
t5.27 stiff and intercostal veins twist
t5.28 Sulphur Component of some amino acids (cysteine Symptoms appear in younger most formed
t5.29 and methionine) and coenzymes leaves, a general chlorosis of entire leaf
t5.30 including vascular bundles
t5.31 Iron Involved in chlorophyll biosynthesis, Symptoms appear first in younger growing
t5.32 structural component of cytochromes and organs, interveinal chlorosis; in severe cases,
t5.33 ferredoxin leaves become white which later dries out
t5.34 Chlorine Stomatal regulation in some plants (e.g. Symptoms appear first in younger leaf,
t5.35 onion), stimulation of proton pumping reduction in leaf surface area, wilting of leaf
t5.36 ATPase located at tonoplast, involved in margins, interveinal chlorosis of mature leaves,
t5.37 photosynthetic oxygen evolution, highly branched root system
t5.38 osmoregulation
t5.39 Copper Activator of some enzymes like Cu-Zn Young leaves turn dark-green colour, twisted
t5.40 SOD with necrotic spots, depressed internode
t5.41 growth, bushy appearance, stunted growth,
t5.42 reduction in panicle formation
t5.43 Manganese Activator of some enzymes (Mn-SOD), Small yellow spots and interveinal chlorosis on
t5.44 involved in PS I as water-splitting complex younger leaves
t5.45 Zinc Activator of some enzymes, involved in Stunted growth due to shortening of internode
t5.46 synthesis of auxin called ‘rosette’, drastic decrease in leaf size
t5.47 Molybdenum Cofactor of enzymes involved in nitrogen Chlorosis and stunted growth in younger leaves,
t5.48 metabolism drastic reduction in size and whiptail
t5.49 appearance of leaf blade
t5.50 Nickel Metal cofactor of urease enzyme Symptoms closely related to Fe deficiency

450 isolated was Chl1 from A. thaliana (renamed as high-affinity components and therefore classified 455
451 AtNRT1.1) primarily expressed in roots, and its as dual-affinity transporter (DATS) (Wang and 456
452 induction was caused by high external nitrate con- Crawford 1996; Wang et al. 1998; Liu et al. 1999). 457
453 centrations up to 25  mM.  Later, kinetic studies Another exception of DATS was reported in 458
[AU3]
454 revealed that AtNRT1.1 possessed both low- and Medicago truncatula, MtNRT1.3 (Morere-Le 459
 20  Mineral Nutrition of Plants

460 Paven et  al. 2011). Very recently, the molecular chlorophyll. Typical symptom of N deficiency is 507
461 mechanism of switching affinity of NRT1.1 from the formation of ‘V’ shape of chlorosis starting 508
462 low to high revealed that it is controlled by phos- from the tip of the leaf (Fig. 20.3). This yellow- 509
463 phorylation and dephosphorylation of a key amino ing later can be seen in younger leaves, and in 510
464 acid residue, threonine, at position 101 (Parker case of severe N deficiency, the older leaves fall 511
465 and Newstead 2014). In Arabidopsis, the entire off. The plant becomes pale green and the leaf 512
466 family of NRT1 and NRT2 genes have been char- petiole and veins become purple due to anthocy- 513
467 acterised comprising of nine and seven members, anin pigment synthesis. The overall plant growth 514
468 respectively, that control the flux of nitrate from is poor and stunted due to low protein synthesis. 515
469 soil into root tissues and throughout the plant The symptom appears later in the young leaves 516
470 body. These transport systems are induced by because N is highly mobile in plant, and so it is 517
471 NO3− concentration in soil solution as well as the translocated from older leaves to the young 518
472 N status of the plant. Besides nitrate transporters, growing points. 519
473 a large number of high- and low-affinity ammo- Plants grown with excessive N usually have 520
474 nium transporters are encoded by the AMT family dark-green-coloured leaves, produce excess foli- 521
475 in plants grown under reduced soil conditions age but have a poorly developed root system and, 522
476 (Howitt and Udvardi 2000). Organic N forms are thus, a low root/shoot ratio. For example, potato 523
477 also transported throughout the plant by proton- plants supplied with excess N produce profuse 524
478 dependent oligopeptide transporters of the POT/ foliage with a few small tubers and poor root 525
479 PTR family. growth; perhaps sugar translocation to tubers and 526
root is affected due to hormonal imbalance. 527
480 20.4.1.1.2  A  ssimilation and Biological Flowering and seed development in several agri- 528
481 Functions of N cultural and horticultural crops are reduced; the 529
482 Plants cannot use inorganic N as such so it has to flowering is delayed due to excessive vegetative 530
483 be reduced. Two important enzymes of N assimi- growth. However, short-day plants give abundant 531
484 lation, nitrate reductase and nitrite reductase, are N flower faster. Excess N also causes tomato 532
485 involved in reducing the oxidised form of N, that fruits to split as they ripen. 533
486 is, NO3− to NH4+. Other enzymes of N assimila-
487 tion pathway include glutamine synthetase (GS), 20.4.1.2 Phosphorus 534
488 glutamate synthase (GOGAT), glutamate dehy- Almost 90 % of P is fixed in soil in the form of 535
489 drogenase (GDH), aspartate aminotransferase aluminium/iron phosphates or c­ alcium/magne- 536
490 (AspAT) and asparagine synthetase (AS). These sium phosphates depending on soil pH. Plants 537
491 enzymes are responsible for the incorporation of cannot use these fixed or non-labile forms of 538
492 NH4+ into amino acids such as glutamine, gluta- P. Another part of insoluble P, called the labile 539
493 mate, asparagine and aspartate. The primary fraction, exchanges with the soil solution. The 540
[AU4]
494 function of N is to provide amino groups in inorganic P released from the labile fraction into 541
495 amino acid constituent of bases in nucleotides of the soil solution, called the solution P, is 542
496 purine and pyrimidine. Besides these, N is an extremely slow and can take a few years. This is 543
497 essential constituent of many nonprotein com- the only form of P accessible to plants for 544
498 pounds such as coenzymes, photosynthetic pig- uptake. Therefore, P deficiency is a widespread 545
499 ments, secondary metabolites and polyamines phenomenon. Since phosphatic fertilisers are 546
500 and vitamins. When N is in ample supply, the made from rock phosphates, P is considered as a 547
501 NO3− form is stored in the vacuole where it con- nonrenewable resource which is expected to be 548
502 tributes to generation of turgor pressure. exhausted within the next 50–60  years. The 549
form in which P is found in soil solution is pH 550
503 20.4.1.1.3  S  ymptoms of Deficiency dependent (Fig. 20.1), but at typical soil solu- 551
504 and Excess N tion pH, P occurs exclusively as H2PO4−, the 552
505 Deficiency symptom is seen as a general chloro- preferred form of inorganic P (Pi) taken up by 553
506 sis starting in the lower leaves due to loss of plants. 554

Fig. 20.3  Deficiency symptoms of macronutrients cre-
ated in maize seedlings (15 days old) grown in hydro-
ponics. The element in question was omitted from the
nutrient solution thus producing severe deficiency
symptoms. –N leaf showing characteristic ‘V’ shape
chlorosis starting from tip of leaf, stubby root growth
without laterals. –P shoot growth more depressed than
555 20.4.1.2.1  P Uptake and Distribution
556 In response to persistent Pi deficiency, plants
557 have developed many adaptive morphological,
558 physiological and molecular mechanisms to cope
559 with low Pi. These includes changes in root
560 growth (increased root surface area, fine root
561 hairs, root length) and architecture, induction of
562 high-affinity Pi transporters, increased secretion
563 of acid phosphatase enzyme and low-molecular-­
564 weight organic acids, symbiotic associations with
565 mycorrhizal fungi and changes in the activity of
R. Pandey
root growth, dark-green leaves. –K leaves with marginal
necrosis, poor root growth without lateral roots. –Ca
characteristic symptom is death of growing point, bushy
roots. –Mg interveinal chlorosis in older leaves, purple
coloration due to anthocyanin. –S interveinal chlorosis
in younger leaves (Symptoms developed by Renu
Pandey)
several key photosynthetic enzymes (reviewed by 566
Lopez-Arredondo et al. 2014). Several plant spe- 567
cies (e.g. Proteaceae) form dense clusters of fine 568
lateral roots called proteoid roots under P defi- 569
ciency. These clusters release large amounts of 570
organic acids that act as chelators and helps in 571
bringing sparingly soluble calcium phosphates 572
into soil solution. 573
A wide variety of Pi transporter family has 574
been identified in plants recently based on genome 575
sequence analysis and experimental evidences. 576
 20  Mineral Nutrition of Plants

577 These transporters are involved in the uptake of to hydroxyl groups to a carbon chain (C-O-P) as 609
578 Pi, movement within the cell and around the plant a simple phosphate ester (e.g. sugar phosphate) 610
579 body. Based on their protein sequence, structure, or attached to another phosphate by the energy-­ 611
580 localisation in membrane and functions, they have rich pyrophosphate bond (P  ~ P), such as in 612
581 been grouped into different families. These ATP. Another type of phosphate bond is the dies- 613
582 include Pht1 (plasma membrane), Pht2 (plastid ter state (C-P-C) with relatively high stability. In 614
583 inner envelope), Pht3 (mitochondrial inner mem- this association, phosphate forms a bridging 615
584 brane), Pht4 (chloroplasts, heterotrophic plastids group between connecting units resulting in more 616
585 and Golgi) and pPT (plastid inner envelope). complex macromolecular structures. P as a struc- 617
586 Among the Pi transporters, Pht1 family is most tural element is a constituent of nucleic acids 618
587 widely studied which belongs to high-affinity Pi (DNA, RNA) (responsible for their strongly 619
588 transporter family involved in Pi uptake by roots. acidic nature) and phospholipids of biomem- 620
589 High-affinity Pi transporter operates at low exter- branes forming phosphatidylcholine (lecithin). In 621
590 nal Pi concentrations (l–10 μM), whereas the low- membranes, Pi acts as a link between glycerol-­ 622
591 affinity Pi transporter, which is a Pi/H+ symporter, fatty acid (lipophilic part) and the choline (hydro- 623
592 has an apparent Km of 0.4  mM (Rausch and philic) part of the lipid. Choline is strongly 624
593 Bucher 2002). The Pht1 families from Arabidopsis hydrophilic because of the negative charge on 625
594 and rice contain 9 and 13 members, respectively, phosphate group, and this helps in proper orienta- 626
595 and have been well characterised (Mudge et  al. tion in the membrane. Further, P has a major role 627
596 2002). The presence of mycorrhizal associations in energy transfer reactions. This involves forma- 628
597 also influences the expression of specific trans- tion and disruption of pyrophosphate bond to 629
598 porters. Vacuolar sequestration of P occurs during maintain energy homeostasis in cellular pro- 630
599 seed development where large amounts of P and cesses. Hydrolysis of one mole of ATP releases 631
600 other minerals (Ca, Mg, Zn, Fe) are stored in 30  kJ of energy. ATP is the basis of many syn- 632
601 seeds forming a complex inositol-hexaphosphate thetic pathways, and other similar energy-rich 633
602 called phytate. phosphonucleotides (UTP, CTP and GTP) play a 634
central role in nucleic acid metabolism. UTP is 635
603 20.4.1.2.2  A ssimilation and Biological involved in synthesis of sucrose, starch and cel- 636
604 Functions of P lulose, while CTP provides energy during phos- 637
605 Unlike N and S, P remains in its highly oxidised pholipid biosynthesis. 638
606 form once it enters the plant. The inorganic P is The energy-rich phosphates like ATP, GTP or 639
607 either found as soluble Pi (orthophosphate) or as ADP modulate enzyme activities by reversible 640
608 PPi (pyrophosphate). Organic P is mainly bound phosphorylation (Fig. 20.4). This regulatory 641

Fig. 20.4  Modulation X~P X

of enzyme activity by
reversible protein Ca2+ (Calmodulin)
phosphorylation
Protein kinase

Enzyme inactive Enzyme – Pactive

Protein phosphatase
X = ATP

Pi H2O
 R. Pandey

642 phosphorylation is mediated by protein kinases Because P is highly mobile inside the growing 689
643 and can result in activation, inactivation and/or tissues in plants, the older leaves are first to show 690
644 changes in the allosteric properties of the target chlorosis. Plant maturity is also delayed. 691
645 molecule. Protein kinase phosphorylates serine Excess P application leads to P toxicity in 692
646 or threonine residues of proteins and the enzyme plants leading to delayed formation of reproduc- 693
647 becomes activated, while dephosphorylation is tive organs. This may be because P and N have 694
648 carried out by phosphatases that release Pi, hence synergistic effect, meaning presence of P will 695
649 inactivating the enzyme protein. Protein phos- lead to more uptake of N. 696
650 phorylation also has an important role in signal
651 transduction. 20.4.1.3 Potassium 697
652 The large amounts of P stored in seeds as The Earth’s crust contains around 2.3  % 698
653 phytic acid facilitate development of embryo, K.  Mostly K is bound in primary minerals or 699
654 seed germination and seedling growth. The con- present in secondary clay minerals making clayey 700
655 centration of Pi has a very important role in the soils rich in K. Examples of some of the minerals 701
656 process of photosynthesis. Generally photosyn- containing K as K2O are alkali feldspar (4–15 %), 702
657 thesis is limited by Rubisco activity or the capac- muscovite or K mica (7–11  %), biotite or Mg 703
658 ity to regenerate ribulose 1,5-bisphosphate mica (6–10 %) and illite (4–7 %). A typical con- 704
659 (RuBP). In light, for optimal photosynthesis, a Pi centration of K in soil solution varies between 0.1 705
660 concentration in the range of 2.0–2.5  mM is and 1  mM K+. Soil K occurs in three forms: K 706
661 required in the chloroplast; however, photosyn- present in soil solution (readily available to 707
662 thesis is inhibited if the Pi concentration falls plant), K adsorbed in exchangeable form to soil 708
663 below 1.4–1.0  mM (Marschner and Marschner colloids such as clay minerals and K as a struc- 709
664 2012). This may be due to many of the intermedi- tural element of soil minerals. Generally, K defi- 710
665 ary steps involving sugar phosphates during car- ciency is a rare occurrence, but plant growth is 711
666 bon fixation. Therefore, the demand for Pi is usually stimulated by additional K supply. 712
667 higher in chloroplast which is achieved by the Pi/
668 triose phosphate (Pi/TP) translocator present in 20.4.1.3.1  K Uptake and Distribution 713
669 the chloroplast envelope. The synthesis of starch Plants take up potassium as monovalent cation, 714
670 in chloroplast is regulated by Pi/TP ratio. A high K+. Uptake of K in plant tissues occurs at high 715
671 Pi/TP ratio in chloroplast inhibits the key enzyme rates due to relatively high permeability of plant 716
672 ADP-glucose pyrophosphorylase, thus decreas- membranes to K. This high permeability of mem- 717
673 ing starch synthesis. However, a low Pi/TP ratio branes to K results from ionophores located in 718
674 activates this enzyme, and TPs are diverted membrane that enables facilitated diffusion. 719
675 towards sucrose synthesis in the cytoplasm. Besides the passive uptake, K also enters plant 720
roots via high- and low-affinity transporters 721
676 20.4.1.2.3  S  ymptoms of Deficiency (Giertha and Maser 2007; Wang and Wu 2013). 722
677 and Excess P Studies employing electrophysiology techniques 723
678 Deficiency of P causes stunted growth of plants indicated that passive transport of K occurs 724
679 and foliage is often dark green. The dark-green through ion channels with millimolar Km denot- 725
680 colour of leaves is because of the accumulation ing low-affinity and active transport through H+- 726
681 of starch and sugars in the leaves. Under P defi- cotransporters with micromolar Km denoting 727
682 ciency, shoot growth is much more repressed high-affinity transporters. Once inside the plant, 728
683 than root growth leading to a higher root/shoot K is highly mobile at all levels, i.e. within indi- 729
684 ratio (Fig. 20.3). In severe cases, the roots also vidual cells, tissues and in long-distance trans- 730
685 develop purple coloration (observed in hydro- port via the xylem and phloem. The bulk of K is 731
686 ponically grown maize plants). The leaf expan- taken up during vegetative phase. A large flux of 732
687 sion is affected due to reduced cell division and K from shoot to root is maintained through 733
688 enlargement thereby producing smaller leaves. phloem, which is crucial to maintain K homeo- 734
 20  Mineral Nutrition of Plants

735 stasis and to provide a constant supply of cations ute transport causes cell extension, stomatal 782
736 to accompany anions like NO3− for their move- movements and other photonastic and seismo- 783
737 ment towards the shoot. nastic movements. Moreover, the loading and 784
unloading of sucrose in phloem are also depen- 785
738 20.4.1.3.2  Biological Functions of K dent on K concentration in the sieve tubes. 786
739 Unlike other elements, K is not metabolised in the
740 plant and it forms only weak complexes in which 20.4.1.3.3  S ymptoms of Deficiency 787
741 it is readily exchangeable. K has an exceptional and Excess K 788
742 role in plant-water relations. Besides maintaining As with N and P, K is also easily redistributed in 789
743 turgor, it is required for activating an array of plant tissue, so the deficiency symptoms first 790
744 enzymes in metabolic reactions. It is maintained appear on the older leaves. The typical symptom 791
745 in the range of 100–200 mM in the cytosol, and of K deficiency is the development of chlorosis 792
746 almost similar concentration is found in chloro- which later turns into necrotic lesions on the leaf 793
747 plast. This storage pool is called ‘metabolic pool’, tip spreading downwards on the margins. Under 794
748 which is not replaceable by other inorganic cat- severe K deficiency, the younger leaves also 795
749 ions such as Na+. However, K concentration in become chlorotic (Fig. 20.3). In maize, root 796
750 vacuole may vary between 10 and 200  mM or development is poor without lateral roots and the 797
751 sometimes may even reach 500 mM, and this stor- stalks are weak. Lignification of vascular bundle 798
752 age pool is called ‘non-­ metabolic pool’, fre- is generally impaired making the plant prone to 799
753 quently replaced by other cations. lodging. Other characters of K deficiency are 800
754 In cytosol, K is involved in enzyme activation; rolling of leaves and shortening of internodes 801
755 specific enzymes include vacuolar pyrophospha- leading to stunted growth. Severe K deficiency 802
756 tases (PPases) that accumulate protons into the causes accumulation of reducing sugars and 803
757 vacuolar lumen and are strictly dependent on depletion of organic acids and synthesis of toxic 804
758 K. Besides this, many enzymes involved in car- amines such as putrescine and agmatine by the 805
759 bon metabolism such as pyruvate kinase, phos- decarboxylation of arginine. 806
760 phofructokinase and ADP-glucose starch High K content in the growing medium does 807
761 synthase are also activated by K. It is essential for not usually produce any toxic symptoms in plant. 808
762 chlorophyll development and catalyses normal This may be because uptake of K in excess repre- 809
763 carbohydrate breakdown during respiration. sents ‘luxury consumption’ and it is stored in the 810
764 Protein synthesis mediated by ribosomes is vacuole without any increment in growth. 811
765 another key process that requires high concentra-
766 tions of K. The metabolic enzymes involved in 20.4.1.4 Sulphur 812
767 transcriptional and post-transcriptional regula- Soils contain inorganic and organic forms of S, 813
768 tion are also affected by K status of the plants, but the major soil S reservoir is organically bound 814
769 thereby influencing the metabolism. K is involved S. The organic S is mostly present in the form of 815
770 in the up-regulation of malic enzyme and assimi- phenolic and choline sulphates as well as lipids 816
771 lation of nitrate via the GS/GOGAT pathway, and amino acids, and the C/N/S ratio in soil 817
772 while the uptake of nitrate and its reduction are organic matter is approximately 125:10:1.2. In 818
773 downregulated (Armengaud et al. 2009). It plays saline and sodic soils, inorganic salts are predom- 819
774 an important role in photosynthesis since it is the inant. Under aerobic soil condition, inorganic S is 820
775 dominant counterion to the light-induced proton present primarily as sulphate (SO42−): the pre- 821
776 flux across the thylakoid membrane and for ferred form for uptake by plants. Under water- 822
777 establishing the transmembrane pH gradient logged conditions, inorganic S occurs in reduced 823
778 required for ATP synthesis (photophosphoryla- forms such as FeS, FeS2 and H2S. Sulphate reduc- 824
779 tion). The dominant role of K is in turgor mainte- tion under waterlogged (anaerobic) condition is 825
780 nance and water homeostasis termed as carried out by bacteria belonging to genus 826
781 osmoregulation. The turgor pressure-driven sol- Desulfovibrio leading to formation of H2S. The 827
 R. Pandey

828 H2S undergoes oxidation to elemental S by presence of light, while some may be reduced in 875
829 c­ hemotrophic S bacteria such as Beggiatoa and root plastids. Surplus S is deposited in vacuoles as 876
830 Thiothrix. The total S content in soil varies in the SO42−. SO2 when reacts with water in cells forms 877
831 range from 0.005 to 0.04 %. bisulphite (HSO3−), and in this form, it inhibits pho- 878
tosynthesis and causes chlorophyll destruction. 879
832 20.4.1.4.1  S  Uptake, Distribution S assimilation involves activation of SO42− 880
833 and Assimilation ions by ATP forming adenosine phosphosulphate 881
834 In addition to S uptake by roots as SO42−, plants (APS). The APS then serves as a substrate for the 882
835 can also absorb S from the atmosphere in the synthesis of sulphate esters (sulpholipids), poly- 883
836 form of SO2 through stomatal openings. Uptake saccharides or secondary metabolite (glucosino- 884
837 of SO42− is an active process as it is absorbed via lates). Through another pathway, APS is reduced 885
838 plasma membrane sulphate transporters ener- and incorporated into amino acid cysteine, the 886
839 gised by H+ gradient. The proton-coupled first stable product of a series of reactions involv- 887
840 cotransport occurs with 3H+/SO42− stoichiometry. ing glutathione, reduced ferredoxin and acetyl 888
841 Transcription of this type of mechanism is serine. The enzymes of assimilatory SO42− reduc- 889
842 induced when S becomes deficient. Analysis of tion are localised in the chloroplasts and to some 890
843 genome sequences of Arabidopsis and rice has extent also found in roots. 891
844 led to the identification of 14 putative sulphate
845 transporter genes in each genome. These genes 20.4.1.4.2  Biological Functions of S 892
846 are subdivided into 4 closely related groups and Most S is found in reduced form in the amino 893
847 all having 12 membrane-spanning domains and a acids cysteine and methionine and hence a con- 894
848 STAS domain at their carboxy-terminus (Aravind stituent of protein. Both of these amino acids are 895
849 and Koonin 2000). The fifth group is more diverse precursors of other S-containing compounds 896
850 but related with two smaller proteins lacking the such as coenzymes (CoA), vitamins (biotin, thia- 897
851 STAS domain (Hawkesford 2003). However, mine) and secondary metabolite (glucosinolates, 898
852 only groups 1 and 2 transporters have sulphate alliins). S is a structural constituent of these 899
853 transport activity. In roots, the genes belonging to ­compounds (e.g. R1-C-S-C-R2) or acts as a func- 900
854 Sultr family are involved in sulphate uptakes that tional group, sulfhydryl (R-SH or thiol), that can 901
855 are located in epidermal and cortical plasma undergo reversible oxidation. The formation and 902
856 membranes. The high-affinity sulphate transport- disruption of these S bridges affect the tertiary 903
857 ers which belong to group 1 operate at low sul- and quaternary structure of protein, thereby influ- 904
858 phur concentration (Km: 1.5–10  μM). However, encing protein activity. The production of gluta- 905
859 group 2 sulphate transporters belong to low-­ thione, a tripeptide having –SH group, serves 906
860 affinity transport activity and are constitutively many functions in plant. Glutathione is readily 907
861 expressed under high sulphate concentration (Km: water soluble and acts as a powerful antioxidant, 908
862 99.2  μM and 1.2  mM). Group 2 transporters, present mostly in chloroplasts. Glutathione may 909
863 Sultr2;1 (Km 0.41  mM) and Sultr2;2 (Km function as a transient storage pool of reduced S 910
864 1.2 mM), have specific functions in the vascular and also a precursor of phytochelatins or ‘class 911
865 movement of sulphate. Sultr2;1 is expressed in III metallothioneins’. These phytochelatins have 912
866 the xylem parenchyma and phloem cells in a general structure of (Glu-Cys)n-Gly, where n is 913
867 leaves, but Sultr2;2 is localised specifically in between two and more than ten and functions in 914
868 phloem of roots and vascular bundle sheath cells detoxifying heavy metals. Research in the direc- 915
869 of leaves (Buchner et al. 2004). tion of increasing phytochelatin production is 916
870 Inside plants, SO42− is highly mobile resulting in being carried out in order to make (crop) plants 917
871 its rapid transport from root to shoot tissues through more tolerant to pollutants such as arsenic and 918
872 xylem. Its translocation is mainly in an upward Cd2+ and to improve the phytoremediation poten- 919
873 (acropetal) direction. Once sulphate is taken up, tial. Another important family of thiols in plants 920
874 bulk of it is reduced in shoot chloroplasts in the is thioredoxins, which functions as regulatory 921
 20  Mineral Nutrition of Plants

922 proteins in carbon metabolism. In reduced form, are well enough saturated with Ca2+ to adequately 968
923 thioredoxin activates several enzymes of Calvin meet the crop demand. 969
924 cycle.
925 The biomembranes contain S as sulpholipids; 20.4.1.5.1  Ca2+ Uptake and Distribution 970
926 in chloroplast thylakoids, they are essential for Plants absorb calcium as a divalent cation, Ca2+. 971
927 the stabilisation of photosystem components. The uptake potential of Ca is lower than other 972
928 Sulpholipids are also involved in the regulation cations such as K+ even though the Ca concentra- 973
929 of ion transport across biomembranes. High lev- tion of soil solution is ten times higher than 974
930 els of sulpholipids in roots provide tolerance to K. This is because Ca absorption takes place only 975
931 plants against salt. S in glucosinolates is stored in through young root tips in which the endodermis 976
932 vacuoles, and their hydrolysis catalysed by cell walls are still unsuberised. Uptake of Ca 977
933 enzyme myrosinase releases sulphate that can be from soil solution by roots is mainly a passive 978
934 recycled under S-deficient condition (Grubb and process, and even within the plant tissues, Ca 979
935 Abel 2006). translocation occurs passively. The upward trans- 980
location of Ca in xylem sap occurs with the tran- 981
936 20.4.1.4.3  S  ymptoms of Deficiency spiration stream. Since Ca precipitates as 982
937 and Excess S Ca-phosphate in the phloem sap, the downward 983
938 Unlike nitrogen, S deficiency symptom first translocation of Ca is very slow. Ca enters the 984
939 appears in the younger most leaf (Fig. 20.3). root through Ca2+-permeable channels. Some of 985
940 Chlorosis occurs throughout the entire leaf includ- these channels are Ca2+ selective, but others are 986
941 ing vascular bundles (veins). Under severe S defi- ‘nonselective’ ion channels. The identity of the 987
942 ciency, inhibition of protein synthesis takes place specific protein(s) that mediates Ca2+ uptake is 988
943 that also drastically decreases chlorophyll content still unknown (Demidchik and Maathuis 2007). 989
944 of leaves. In S-deficient plants, there is an accu- Within the plant, the major Ca2+ transporter at the 990
945 mulation of soluble organic nitrogen and nitrate. plasma membrane and also at endoplasmic 991
946 Plants are comparatively insensitive to high ­reticulum is a Ca-pumping ATPase (Ca2+/H+ anti- 992
947 SO42− concentrations in the nutrient media. porter). However, inside the plant, Ca is rather 993
948 However, if the SO42− concentration increases immobile; the ions have a tendency to be seques- 994
949 beyond 50  mM as in some saline soils, plant tered in the large vacuole of mature cells. No 995
950 growth is adversely affected. The symptom of transporters have been identified that are respon- 996
951 excessive S is a reduction in growth rate and dark- sible for loading Ca2+ into xylem vessels. A pro- 997
952 green leaves. High SO2 in the environment is toxic portion of xylem Ca2+ is contributed by the 998
953 to plants as it causes bleaching of chlorophyll apoplast. However, Ca2+ mobility is low in the 999
954 characterised by necrotic symptoms in leaves. vascular system. 1000

955 20.4.1.5 Calcium 20.4.1.5.2  Biological Functions of Ca 1001

956 Calcium is abundant in the lithosphere, constitut- Cellular functions of Ca2+ are mainly concerned 1002
957 ing about 3.64  % of the Earth’s crust. The with structure and as a secondary messenger. It is 1003
958 Ca-containing minerals are Al-silicates (feld- a constituent of middle lamella in the form of cal- 1004
959 spars and amphiboles), Ca phosphates and Ca cium pectate and helps to cement the wall of cells 1005
960 carbonates. Severely weathered soils followed by together. The proportion of calcium pectate in the 1006
961 leaching lead to Ca-deficient soil, which is accel- cell walls is of particular importance as it is 1007
962 erated by low soil pH. The Ca2+ adsorbed on soil responsible for the susceptibility of tissue to fun- 1008
963 colloids may be exchanged with the soil solution gal and bacterial infections and also for fruit rip- 1009
964 resulting in ‘free’ Ca2+ that forms insoluble com- ening. Ca is also present as insoluble crystals of 1010
965 pounds with elements such as P, thus making it Ca-oxalate called raphides and sphaeraphides in 1011
966 less available. Most soils contain enough levels the vacuoles of some plant species (e.g. in 1012
967 of Ca2+ in soil solution, and their exchange sites Colocasia, raphides are responsible for itching 1013
 R. Pandey

1014 sensation). Ca is essential for the formation of below a critical level leading to development of 1061
1015 cell membranes and lipid structures. Most of the diseases such as ‘blossom-end rot’ in tomatoes, 1062
1016 Ca present in plant tissues is localised in the apo- ‘black heart’ in celery and ‘bitter pit’ in apples. 1063
1017 plast and in vacuoles. The Ca2+ concentration of Ca influences the permeability of plasma mem- 1064
1018 cytoplasm is low ranging from 10−6 to brane, thus its deficiency makes the membrane 1065
1019 10−8 M. Such a low cytosolic concentration is of leaky. Ca also mediates starch hydrolysis to 1066
1020 vital importance because higher Ca2+ concentra- sugar, hence its deficiency results in the accumu- 1067
1021 tion inhibits various enzymes located in cyto- lation of starch in leaves. The number of mito- 1068
1022 plasm as well as in chloroplasts. However, Ca chondria in wheat roots reduces under Ca 1069
1023 concentration is much higher in the mitochon- deficiency. 1070
1024 dria. In cytosol, Ca is reversibly bound to a small
1025 protein, a polypeptide of 148 amino acids, called 20.4.1.6 Magnesium 1071
1026 calmodulin (calcium-modulated protein). Magnesium is derived from the Greek word 1072
1027 Calmodulin then activates downstream events in ‘Magnesia’. The Mg content of soil varies 1073
1028 the signalling cascade by phosphorylation of sol- between 0.05 and 0.5 %. Easily weatherable fer- 1074
1029 uble and membrane-bound proteins. Ca in small romagnesian minerals such as biotite, serpentine, 1075
1030 amounts is essential for normal mitosis and is hornblende and olivine and other secondary clay 1076
1031 associated with chromatin or mitotic spindle minerals add Mg to the soil. In soil solution just 1077
1032 organisation. like Ca2+, Mg2+ is also present in fairly high con- 1078
1033 The formation of secretory vesicles and their centration between 2 and 5  mM.  The size of 1079
1034 fusion with plasma membrane leading to exocy- hydrated ion is small (0.428  nm radius), due to 1080
1035 tosis require Ca. This helps in the formation of which Mg adsorption to soil particles is relatively 1081
1036 cell wall from the precursor cellulose as well as weak resulting in high leaching loss to the tune of 1082
1037 the formation of mucilage and callose. In root- 2–30 kg per hectare. Such losses lead to recurring 1083
1038 caps, secretion of mucilage depends on extracel- Mg deficiency in crops. 1084
1039 lular (apoplasmic) Ca concentration. One of the
1040 vital functions performed by Ca is the cell mem- 20.4.1.6.1  Mg Uptake and Distribution 1085
1041 brane stabilisation which is brought about by In plants, the free cytoplasmic Mg2+ remains at a 1086
1042 bridging phosphate and carboxylate groups of concentration of about 0.5  mM (Yazaki et  al. 1087
1043 phospholipids and protein at membrane surfaces. 1998), but total Mg2+ levels may vary from 0.3 to 1088
1044 Ca present in vacuole contributes to cation-anion 1.0 %. Transport of Mg2+ is passive, mediated by 1089
1045 balance by acting as a counterion for inorganic ionophores in which Mg2+ moves down against 1090
1046 and organic anions. Free Ca at a very low cyto- the electrochemical gradient. In this transport 1091
1047 solic concentration (0.1–0.2 μM) acts as second system, Mg2+ faces cation competition; therefore, 1092
1048 messenger. A wide range of stimuli evokes rapid if an excess of other cations such as K+ or NH4+ is 1093
1049 changes in free cytosolic Ca2+ in plants including present in the medium, the uptake of Mg2+ is 1094
1050 responses to abiotic and biotic stresses, stomatal adversely affected. Mg2+ is very mobile in the 1095
1051 regulation and physical damage. phloem and so can be translocated from older to 1096
younger leaves or the meristem. Since Mg2+ has a 1097
1052 20.4.1.5.3  Symptoms of Ca Deficiency primary role in photosynthesis, therefore, highest 1098
1053 Characteristic symptom of severe Ca deficiency tissue Mg2+ concentration is usually measured in 1099
1054 is the twisting and deformation of growing tips shoots. Most of the Mg2+ taken up by plants are 1100
1055 and youngest leaves (Fig. 20.3). At more stored in the vacuole where it contributes to tur- 1101
1056 advanced stage, necrosis occurs at the leaf mar- gor generation and charge balancing of anions. 1102
1057 gin. Ca deficiency also causes disintegration of
1058 cell walls and the collapse of the affected tissues 20.4.1.6.2  Biological Functions of Mg 1103
1059 such as the petioles and upper parts of the stems. Mg occupies the central position in the porphyrin 1104
1060 In fast-growing tissues, Ca2+ levels may fall structure of chlorophyll molecule where it coordi- 1105
 20  Mineral Nutrition of Plants

1106 nates covalently with four N atoms. Insertion of ary structure. Mg stabilises the ribosomal subunits 1135
1107 Mg2+ atom in the porphyrin structure during chlo- necessary for protein synthesis and also has a 1136
1108 rophyll biosynthesis is catalysed by the enzyme similar stabilising effect in the matrix of the 1137
1109 Mg2+-chelatase (Sirijovski et al. 2008). Mg plays nucleus. Further, the transfer of aminoacyls from 1138
1110 a crucial role in photosynthesis, particularly in aminoacyl tRNA to the polypeptide chain is also 1139
1111 promoting the light reactions in the stroma of activated by Mg2+ (Fig. 20.6). [AU5]
1140
1112 chloroplast. Carbohydrate partitioning is also reg-
1113 ulated by Mg2+ concentration in the tissues. The 20.4.1.6.3  Symptoms of Mg Deficiency 1141
1114 loading of sucrose into the phloem in source leaf The first symptom of Mg deficiency is interveinal 1142
1115 requires H+-pumping ATPase, and for the optimal chlorosis of older leaves. In maize seedlings, 1143
1116 activity of this enzyme, about 2  mM Mg2+ is leaves develop purple coloration along with 1144
1117 essential. The majority of cellular Mg2+ functions necrotic spots when Mg was completely omitted 1145
1118 as enzyme cofactors in the stabilisation of nucleo- from the growth medium (Fig. 20.3). In dicotyle- 1146
1119 tides and nucleic acids. Some important enzymes donous plants including grapes, beans, potatoes 1147
1120 activated by Mg2+ are phosphokinases, dehydro- and sugar beet, Mg2+-deficient leaves become 1148
1121 genases and enolases. Activation of ribulose 1,5 stiff and brittle and the intercostal veins twist in 1149
1122 bisphosphate carboxylase during light reaction addition to chlorosis. Protein synthesis is also 1150
1123 occurs due to Mg2+ influx into the thylakoid. The adversely affected resulting in poor growth of 1151
1124 most prominent enzyme reactions where Mg2+ is plants. Other ultrastructural changes in Mg2+- 1152
1125 indispensable are those associated with energy deficient leaves include irregular shape of grana 1153
1126 transfer and phosphorylation/dephosphorylation. with reduction in their numbers, accumulation of 1154
1127 In these reactions, Mg2+ forms a bridge between starch grains in chloroplast, deformation of the 1155
1128 the pyrophosphate structure of ATP or ADP and lamellar structure, underdeveloped cristae of 1156
1129 the enzyme molecule (Fig. 20.5). Mg plays an mitochondria and decrease in chlorophyll and 1157
1130 important role in gene transcription and transla- carotenoid contents. These symptoms of ultra- 1158
1131 tion as it readily binds to nucleic acids. As a result, structural disorganisation lead to visual Mg defi- 1159
1132 DNA-melting temperatures remain considerably ciency symptoms. In maize roots, increased 1160
1133 higher in the presence of Mg2+. In RNA also, Mg2+ suberisation of endodermis and hypodermis 1161
1134 has similar roles and helps in maintaining second- occurs due to Mg deficiency. 1162

Protein
P ~ P ~P
N
Mg ..
Mg
N N N N
Adenine Ribose Adenine Ribose P~ P ~ P

Fig. 20.5  Magnesium acts as bridging molecule between nitrogen and phosphoryl groups in ATP (Source: Marschner
1995)

NAPD+ (photosynthesis)
Cys S S S Cys
Nitrite reductase
e- e-
Fe Fe Sulphite reductase
N2 reduction
Cys S S S Cys
GOGAT

Fig. 20.6  Role of ferredoxin as an electron carrier in a number of basic metabolic processes (Source: Marschner 1995)
 R. Pandey

1163 20.4.2 Physiological Functions (Mukherjee et  al. 2006). The mechanism of Fe 1208
1164 of Micronutrients uptake by roots is strongly regulated by a com- 1209
plex system, which involves transcription factor, 1210
1165 20.4.2.1 Iron bHLH, of which PYE and FIT/FER play a central 1211
1166 Iron makes up about 5  % by weight of Earth’s role (Ivanov et al. 2012). Besides IRT, Fe uptake 1212
1167 crust and is present in almost all soils. The primary takes place through other transporters present in 1213
1168 minerals of Fe are ferromagnesian silicates such as the leaf cells. These include several members of 1214
1169 olivine, augite, hornblende and biotite. The con- the YSL (yellow stripe-like) family expressed in 1215
1170 centration of plant available Fe in soils is extremely leaves (such as AtYSL1, AtYSL3 and AtYSL2 in 1216
1171 low. The Fe solubility in soil is largely controlled A thaliana and OsYSL2 and OsYSL15 in rice), 1217
1172 by pH of soil solution. At higher pH (7.4–8.5), the but their expression is usually confined to the 1218
1173 solubility is at minimum, while at low pH or acid, vascular tissue (Curie et al. 2009). 1219
1174 soils Fe availability is very high. Further, in aer- In rapidly growing plants, about 80 % of Fe is 1220
1175 ated soils, maintained at physiological pH range, stored in chloroplasts. It is localised in the plastid 1221
1176 the concentrations of Fe2+ and Fe3+ are below 10−15 stroma as phytoferritin. Some amount of phyto- 1222
1177 M. Under waterlogged or reduced soil condition, ferritin is also detected in xylem, phloem and 1223
1178 Fe3+ is reduced to F2+ and this reduction is brought seeds. It also acts as storage for Fe in nodules of 1224
1179 about by anaerobic bacteria which uses Fe oxides legumes. The major form of Fe transported 1225
1180 as electron acceptors in respiration. through the xylem is ferric citrate. 1226

1181 20.4.2.1.1  Uptake and Distribution 20.4.2.1.2  Biological Functions of Fe 1227

1182 The preferred form of Fe by plants is the reduced Being a redox-active metal, Fe plays a role in 1228
1183 cation Fe2+. Fe cannot be taken up in ionic form photosynthesis, mitochondrial respiration, 1229
1184 as such; it has to chelate with other compounds ­assimilation of nitrogen, synthesis of chlorophyll 1230
1185 that facilitate Fe uptake. Plant species differ in and hormone (ethylene, gibberellic acid, jas- 1231
1186 their ability to utilise sparingly soluble inorganic monic acid), osmoprotection, pathogen defence 1232
1187 Fe and Fe chelates. Plants grown under Fe stress and production and scavenging of reactive oxy- 1233
1188 show physiological and morphological changes gen species. Based on the type of iron ligand, 1234
1189 that help in Fe uptake. The mechanism of Fe there are three groups of Fe-containing proteins: 1235
1190 uptake in plants via roots is based on two strate- 1. Proteins with Fe-sulphur clusters (Fe-S): The 1236
1191 gies: Strategy I involves reduction of Fe3+ to Fe2+ Fe-S clusters are synthesised from inorganic 1237
1192 carried out by membrane-bound enzyme ferric-­ Fe and sulphide. These nonhemeproteins with 1238
1193 chelate reductase (FCR). The reduced form of Fe-S clusters have a key role in electron trans- 1239
1194 iron is transported into root cells through the fer. They constitute part of substrate-binding 1240
1195 metal transporter protein called iron-regulated sites in enzymes, form iron storage moieties, 1241
1196 transporter (IRT). This mechanism operates in are involved in transcriptional or translational 1242
1197 dicotyledonous plants. Strategy II, a chelation-­ regulation, control protein structure in the 1243
1198 based mechanism occurs in Poaceae (grass) fam- vicinity of the cluster and are also involved in 1244
1199 ily where roots release phytosiderophores (PS) disulphide reduction and sulphur donation 1245
1200 which form stable complexes with Fe3+ and is (e.g. in thioredoxins). Therefore, Fe-S pro- 1246
1201 taken up by plant as Fe3+-PS chelate (Romheld teins serve as enzymes, as electron carriers 1247
1202 and Marschner 1986). The gene encoding FCR (e.g. ferredoxin) and as regulatory proteins 1248
1203 enzyme belongs to the FRO family and IRT (e.g. aconitase). 1249
1204 belongs to ZIP family (Wu et al. 2005; Mukherjee 2. Fe-containing hemeproteins: The well-­1250
1205 et al. 2006; Jeong et al. 2008; Guerinot 2000). characterised hemeproteins are the cyto- 1251
1206 In Arabidopsis, FRO2 gene identified in root has chromes involved in electron transfer reactions 1252
1207 been found to encode for Fe3+-chelate reductase of photosynthetic and respiratory systems, 1253
 20  Mineral Nutrition of Plants

1254 which contain a heme Fe-porphyrin complex. 20.4.2.2 Copper 1300

1255 The oxidative enzymes like catalase, peroxi- The soil contains Cu in divalent cation form, 1301
1256 dase, and NADPH oxidase involved in the Cu2+, in the range of 5–50  ppm. However, the 1302
1257 production and scavenging of free radicals concentration of Cu in soil solution is very low 1303
1258 contain hemeprotein. Another very large 10−8 to 60 × 10−8  M.  More than 98  % of Cu is 1304
1259 group of enzymes called cytochrome P450 bound to organic matter in soil, which is an 1305
1260 also contains hemeprotein. Globins such as important factor regulating Cu mobility in soil. 1306
1261 leghemoglobin in nodules of legumes are Cu is absorbed both as divalent cupric (Cu2+) ion 1307
1262 involved in oxygen binding and transport. in aerated soils or monovalent cuprous (Cu1+) 1308
1263 Nitrite reductase and sulphite reductase ions in waterlogged soils. 1309
1264 enzymes involved in N and S assimilation
1265 contain a siroheme and a Fe-S cluster in the 20.4.2.2.1  Uptake and Distribution 1310
1266 enzyme. Hemeproteins are distributed all over Plants take up Cu in very little quantities, so the 1311
1267 the locations in subcellular organelles, for Cu content in plant tissues varies from 2 to 1312
1268 example, cytochrome P450 is localised in 20 ppm. Cu uptake is an active process requiring 1313
1269 endoplasmic reticulum, catalase in peroxi- metabolic energy. Under physiological pH, Cu 1314
1270 somes and other enzymes in the cytoplasm. exists in two oxidation states Cu1+ and Cu2+ and 1315
1271 3. Other Fe proteins: These proteins are grouped can interchange between these forms (monova- 1316
1272 as nonhemeproteins, which bind Fe ions lent copper is unstable). This property of Cu is 1317
1273 directly. Ferritins or phytoferritins are most responsible for its redox function in biochemical 1318
1274 common among this group of proteins. reactions. Cu has high affinity for N atom of 1319
1275 Ferritins control the interaction between Fe amino groups, sulfhydryl groups, carboxylic 1320
1276 homeostasis and oxidative stress in plants. groups and phenolic groups. More than 98–99 % 1321
1277 These are high-molecular-weight 24-mer pro- of Cu is present in these complexed forms in soil 1322
1278 teins that store up to 4,500 Fe atoms in soluble solution and in xylem and phloem sap. Cu ions 1323
1279 and bioavailable form. Mostly nongreen plas- can catalyse the production of free radicals, 1324
1280 tids such as etioplasts and amyloplasts contain which is potentially toxic leading to the damage 1325
1281 ferritins, but they are not found in mature of proteins, DNA and other biomolecules. 1326
1282 chloroplasts (Briat and Lobreaux 1998). Therefore, immediately following Cu uptake, the 1327
metal-scavenging proteins like metallothionein 1328
1283 20.4.2.1.3  S  ymptoms of Deficiency bind majority of Cu ions, thus preventing it from 1329
1284 and Excess Fe accumulating at a toxic level. 1330
1285 Since Fe is relatively immobile inside the plant
1286 tissue, the visual deficiency symptoms first 20.4.2.2.2  Biological Functions of Cu 1331
1287 appear on young growing organs. The young leaf Cu is of utmost importance for life. It is essential 1332
1288 shows interveinal chlorosis, and in severe cases, for photosynthesis and mitochondrial respiration, 1333
1289 the leaf becomes white (due to loss of chloro- carbon and nitrogen metabolism, oxidative stress 1334
1290 phyll) which later dries out. In contrast, the protection and cell wall synthesis. Three different 1335
1291 mature leaves may not show chlorosis at all. Lack types of Cu-protein exist in plants in which Cu is 1336
1292 of Fe inhibits protein synthesis. the metal component. These are: 1337
1293 Excess Fe uptake leads to toxicity, which is a Type 1: Blue Cu-proteins, without oxidase activ- 1338
1294 major problem in wetland rice-cropping systems. In ity that functions in one-electron transfer. 1339
1295 rice, symptoms of Fe toxicity involve development Common example of this is plastocyanin; 1340
1296 of tiny brown spots that later spreads into uniform more than 50 % of Cu localised in chloroplast 1341
1297 brown colour, called bronzing. The Fe concentra- remains in this form. Plastocyanin is a compo- 1342
1298 tion in rice leaves remains excessively high in the nent of electron transport chain of photosys- 1343
1299 range of 300–1,000 μg Fe per g dry weight. tem I in photosynthesis. 1344
 R. Pandey

1345 Type 2: Non-blue Cu-proteins, which are of Cu to displace other metal ions like Fe from 1392
1346 peroxide-­producing oxidases and oxidise physiologically active sites is associated with Cu 1393
1347 monophenols to diphenols. toxicity symptoms. Thus, the Cu toxicity symp- 1394
1348 Type 3: Multi Cu-proteins, contains at least four tom in plants will manifest as Fe deficiency 1395
1349 Cu atoms per molecule which act as oxidases. symptom. Other symptoms of toxicity include 1396
1350 Multi-Cu enzymes contain all three types of inhibition of root growth more than shoot growth. 1397
1351 Cu, e.g. ascorbate peroxidase and diphenol There are some Cu-tolerant species called metal- 1398
1352 oxidase. Cytochrome oxidase is a mixed lophytes, which can tolerate Cu content as high 1399
1353 Cu-Fe protein catalysing the terminal oxida- as 1,000 μg Cu per g dry weight. 1400
1354 tion in mitochondria.
1355 Recent studies have revealed that more than 20.4.2.3 Zinc 1401
1356 100 Arabidopsis proteins are predicted to be The Zn content of the lithosphere is about 80 ppm 1402
1357 complexed with Cu (Kramer and Clemens 2005). and it is usually present in soil in the range of 1403
1358 The high affinity of Cu to dioxygen molecules 10–300 ppm. The ionic radius of Zn2+ is similar 1404
1359 explains the role of Cu as a catalytic metal in to that of Fe2+ and Mg2+ due to which Zn2+ may 1405
1360 many oxidases. Cu metabolism is also intimately replace these elements by isomorphous substitu- 1406
1361 linked to Fe metabolism. Depending on the bio- tion in the mineral structure. Zn interacts with 1407
1362 availability of Cu and Fe, plants possess enzymes organic matter in soil and forms both soluble and 1408
1363 for the alternative use of Cu and Fe, thus catalys- insoluble Zn-organic complexes. The soluble 1409
1364 ing the same biochemical reaction with com- Zn-organic complexes are mainly associated with 1410
1365 pletely different apoproteins. Such reactions amino, organic and fulvic acids, while the inor- 1411
1366 include Cu-nitrite versus heme-nitrite reductase, ganic complexes are derived from humic acids. 1412
1367 Cu/Zn-superoxide dismutase versus Plants absorb Zn in the form of divalent cation. 1413
1368 Fe-superoxide dismutase and cytochrome oxi-
1369 dase versus di-iron oxidase. Cu has also been 20.4.2.3.1  Uptake and Distribution 1414
1370 reported to be a part of the ethylene receptor and Uptake of Zn is metabolically controlled, that is, 1415
1371 is involved in molybdenum cofactor biosynthesis Zn uptake is an active process requiring energy. 1416
1372 (Rodriguez et al. 1999; Kuper et al. 2004). The form in which Zn is translocated from roots to 1417
shoots is not clear. Mobility of Zn inside the plant 1418
1373 20.4.2.2.3  S ymptoms of Deficiency organs is relatively less, so there is accumulation in 1419
1374 and Excess Cu root tissues particularly when Zn concentration in 1420
1375 Visible symptoms of Cu deficiency appear in the media is high. In older leaves, Zn can become 1421
1376 young leaves that often become dark green in very immobile. Much of the Zn absorbed is local- 1422
1377 colour and are twisted exhibiting necrotic spots. ised in the seeds and grains in the protein bodies in 1423
1378 However, in cereals, the leaf tip becomes white the form of globoid crystals. These globoids con- 1424
1379 and the leaves narrow and twisted. The growth of sist mainly of phytate or salts of phytic acid. 1425
1380 internodes is depressed leading to bushy appear- Zn transport via roots takes place through 1426
1381 ance and stunted growth. Pollen grain viability is transporters belonging to ZIP family (ZRT, IRT-­ 1427
1382 affected resulting in reduction in panicle forma- like protein). ZRT1 and ZRT2 (zinc-regulated 1428
1383 tion. Lignin synthesis is impaired due to lack of transporter) are designated as the high- and low-­ 1429
1384 two important Cu-containing enzymes, pheno- affinity Zn transporters, respectively. Till now, 1430
1385 lase and laccase. Of the total Cu concentration in over 25 ZIP family members have been identified 1431
1386 plants, almost half is found in the chloroplasts which is subdivided into two subfamilies 1432
1387 where it plays an important role in photosynthetic (reviewed by Guerinot 2000). 1433
1388 reactions. Cu deficiency symptom in citrus is
1389 called ‘die back’ because young leaves die out. 20.4.2.3.2  Biological Functions of Zn 1434
1390 The critical toxicity level of Cu in the leaves is Zn has a strong tendency to form tetrahedral 1435
1391 above 20–30 μg Cu per g dry weight. The ability complexes with N-, O- and S- ligands, which 1436
 20  Mineral Nutrition of Plants

1437 help in regulating metabolic functions. Therefore, 3–4  % B.  Only the monomeric species B(OH)3 1483
1438 Zn plays a catalytic (functional) and structural and B(OH)4− are present in soil solution depend- 1484
1439 role in enzyme reactions. Since it exists only as ing on soil pH. 1485
1440 Zn(II), so it does not take part in oxidation-­
1441 reduction reactions. Zn plays a crucial role in 20.4.2.4.1  Uptake and Distribution 1486
1442 many biological processes. It is an integral com- Uptake of B is a non-metabolic process and its 1487
1443 ponent of many enzymes, alcohol dehydroge- distribution in the plants is also governed by tran- 1488
1444 nase, carbonic anhydrase, Cu-Zn-superoxide spiration stream. Boric acid channels, which are 1489
1445 dismutase, alkaline phosphatase, phospholipase, major intrinsic proteins, facilitate B transport 1490
1446 carboxypeptidase and RNA polymerase, to name across membranes. Nodulin26-like intrinsic pro- 1491
1447 a few. Large number of enzymes requires Zn for tein 5;1 (NIP5;1), identified from Arabidopsis, is 1492
1448 activation such as dehydrogenases, aldolases, involved in efficient uptake of B into root cells 1493
1449 isomerases and transphosphorylases. Zinc is under B-stress condition. Another channel pro- 1494
1450 important in DNA and RNA metabolism and pro- tein, AtNIP6;1, helps in preferential distribution 1495
1451 tein synthesis and maintains the structural integ- of B in young shoot tissues (Takano et al. 2006). 1496
1452 rity of biomembranes. More than 1,200 protein Besides boric acid channels, transporters have 1497
1453 molecules (Zn metalloprotein) have been identi- also been identified from Arabidopsis, viz., 1498
1454 fied including a large number of ‘zinc-finger’- BOR1, BOR2 and BOR4 (Takano et  al. 2002). 1499
1455 containing proteins and transcription factors, BOR1 is the first B transporter identified by anal- 1500
1456 oxidoreductases and hydrolytic enzymes such as ysis of Arabidopsis bor1-1 mutant which requires 1501
1457 metalloproteases. Zn is a structural component of high levels of B for normal leaf expansion 1502
1458 ribosomes and thus essential for their structural (30 mM) and fertility (100 mM). BOR1 encodes 1503
1459 integrity. It plays a major role in carbohydrate an efflux-type B transporter which is located in 1504
1460 metabolism by regulating key enzymes, fructose the plasma membrane and is expressed in root 1505
1461 1,6-bisphosphatase and aldolase. Synthesis of cells including endodermis (Noguchi et al. 1997). 1506
1462 auxin, indole acetic acid, is particularly impaired Thus, BOR1 is required for effective xylem load- 1507
1463 under Zn deficiency. Further, Zn also has a role in ing under B-limited conditions, and it is also 1508
1464 signal transduction through mitogen-activated involved in the preferential distribution of B to 1509
1465 protein kinases (MAPK). young leaves. BOR2 encodes a B-efflux trans- 1510
porter located in plasma membrane and strongly 1511
1466 20.4.2.3.3  S  ymptoms of Deficiency expressed in epidermis of elongation zones of 1512
1467 and Excess Zn roots and lateral rootcaps. At toxic B concentra- 1513
1468 The most striking symptoms of Zn deficiency in tions, BOR4 is stably accumulated in plasma 1514
1469 dicotyledonous plants are stunted growth due to membrane and confers high B tolerance to plants 1515
1470 shortening of internodes called ‘rosette’ and a dras- suggesting its involvement in B tolerance by 1516
1471 tic decrease in leaf size termed as ‘little leaf’. When exporting the mineral out of symplast (refer to 1517
1472 the Zn deficiency is severe, the growing shoot apex Miwa and Fujiwara 2010). Certain plant organs 1518
1473 dies, ‘die back’ symptom. Further, these symptoms such as anthers, stigma and ovary contain high 1519
1474 combined with chlorosis produce ‘mottled leaf’. concentrations of B, which is twice as high as in 1520
1475 The critical toxicity levels of Zn in the leaves stem. 1521
1476 ranges between 100 and more than 300 μg Zn per
1477 gram dry weight. Symptom of Zn toxicity is inhi- 20.4.2.4.2  Biological Functions of B 1522
1478 bition of root elongation, chlorosis in young B is involved in a wide range of biological func- 1523
1479 leaves and inhibition of photosynthesis. tions but the exact metabolic functions are not 1524
exactly understood. These important physiological 1525
1480 20.4.2.4 Boron processes include protein synthesis, transport of 1526
1481 The soil contains B in the range of 20–200 ppm. sugars, respiration and the metabolism of plant 1527
1482 The primary mineral tourmaline contains about hormones (indole acetic acid), RNA and carbo- 1528
 R. Pandey

1529 hydrate. Other functions of B are related to cell the divalent cation, Mn2+. Under waterlogged 1575
1530 wall synthesis and lignification, cell wall struc- condition, Mn availability increases just like Fe. 1576
1531 ture maintenance by cross-linking of polysaccha- Mn2+ can replace other divalent cations like Ca2+, 1577
1532 rides and regulation of the structural integrity of Mg2+, Zn2+ and Fe2+ from their active uptake site. 1578
1533 biomembranes. B activates enzymes like plasma-
1534 lemma ATPase thereby increasing the transport 20.4.2.5.1  Uptake and Distribution 1579
1535 of chlorine and phosphorus. Stimulation of H+ Mn uptake is an active process involving meta- 1580
1536 pumping by B causes hyperpolarisation of mem- bolic energy. It is relatively immobile in the plants. 1581
1537 brane potential. Since the wall-associated kinase It is preferentially translocated to meristematic tis- 1582
1538 in the plasma membrane has an extracellular sues; therefore, young plant organs are rich in Mn. 1583
1539 matrix in connection with the pectin molecule, The genes involved in transport of transition metal 1584
1540 the membrane cell wall connection is B depen- in plants have been identified which are also 1585
1541 dent. B promotes structural integrity of biomem- responsible for Mn2+ transport. The gene families 1586
1542 branes and formation of lipid rafts. It also associated with Mn2+ transport include cation/H+ 1587
1543 influences pollen germination, pollen tube growth antiporters, natural resistance-­ associated macro- 1588
1544 and subsequently fertilisation. phage protein (NRAMP) transporters, ZIP trans- 1589
porters, cation diffusion facilitator (CDF) 1590
1545 20.4.2.4.3  S  ymptoms of Deficiency transporter family and P-type ATPases. These 1591
1546 and Excess B transporters are responsible for accumulation of 1592
1547 Death of the root and shoot tips occurs due to B Mn into the cell and its release from various organ- 1593
1548 deficiency resulting in stunted (rosette) plant elles. The active sequestration of Mn into endo- 1594
1549 growth. Leaves develop a thick coppery texture membrane compartments, particularly the vacuole 1595
1550 and become curled and brittle. B also affects and endoplasmic reticulum, also takes place by 1596
1551 flower retention, pollen formation, pollen tube these transporters (reviewed by Pittman 2005). 1597
1552 growth or germination, N fixation and nitrate
1553 assimilation. Root elongation is inhibited and 20.4.2.5.2  Biological Functions of Mn 1598
1554 root tips become swollen and discoloured. The Mn is essential for plant metabolism and approx- 1599
1555 fleshy tissue in fruits disintegrate causing disor- imately 35 enzymes of a plant cell contain Mn in 1600
1556 ders like ‘heart rot’ in sugar beet, ‘water core’ in three oxidation states, II, III and IV. Mn can fulfil 1601
1557 turnip and ‘browning’ of cauliflower. two functions in protein  – it serves as catalyti- 1602
1558 Typical B toxicity symptoms appear on mature cally active metal and it exerts an activating role 1603
1559 leaves producing marginal or tip chlorosis and on enzymes. Enzymes in which Mn has catalytic 1604
1560 necrosis. The critical B toxic content in tissue role are Mn-containing superoxide dismutase 1605
1561 varies depending on the plant species; however, it which protects the cell from damaging effects of 1606
1562 ranges from 100 to 1,000 mg per kg dry weight. free radicals, oxalate oxidase and Mn-containing 1607
water-splitting system of PS II (Barber 2003). 1608
1563 20.4.2.5 Manganese Some of the Mn-activated enzymes are PEP car- 1609
1564 The primary rock minerals containing Mn are boxykinase, isocitrate dehydrogenase, malic 1610
1565 pyrolusite (MnO2) and manganite [MnO (OH)]. enzyme and phenylalanine ammonia lyase (PAL). 1611
1566 Total Mn levels of soil vary between 200 and Manganese activation was seen in enzymes of 1612
1567 3,000 ppm. In biological systems, Mn occurs in nitrogen metabolism (glutamine synthetase, argi- 1613
1568 oxidation states II, III and IV, with Mn(II) and nase), gibberellic acid biosynthesis, RNA poly- 1614
1569 Mn(IV) being fairly stable and Mn(III) unstable. merase activation and fatty acid biosynthesis. 1615
1570 In plants, Mn(II) is the dominant form, but it can
1571 readily undergo oxidation reaction and form 20.4.2.5.3  S
 ymptoms of Deficiency 1616
1572 Mn(III) and Mn(IV). This property makes it pos- and Excess Mn 1617
1573 sible for Mn to play a crucial role in redox reac- Visible symptoms of Mn deficiency first appear 1618
1574 tions. The preferred form of uptake by plants is on younger leaves in the form of small yellow 1619
 20  Mineral Nutrition of Plants

1620 spots and interveinal chlorosis. At ultrastructural 20.4.2.6.2  Biological Functions of Mo 1666
1621 level, Mn deficiency leads to disorganisation of Mo is an essential component (cofactor) of a few 1667
1622 chloroplast and lamellar system. The cell volume important enzymes in higher plants. In these 1668
1623 is reduced, cell walls dominate and the inter-­ enzymes, Mo has both structural and catalytic 1669
1624 epidermal tissue shrinks. ‘Grey speck’ in oats functions and is involved directly in redox reac- 1670
1625 and ‘marsh spot’ in cotyledons of pea are the tions. These enzymes are nitrate reductase, nitro- 1671
1626 symptoms of Mn deficiency. genase, xanthine oxidase/dehydrogenase and 1672
1627 Mn toxicity symptoms are characterised by sulphite reductase. The physiological processes 1673
1628 brown spots in older leaves surrounded by chlo- controlled by these enzymes are N assimilation, 1674
1629 rotic areas. Excess Mn can also induce deficiency S metabolism, phytohormone biosynthesis and 1675
1630 of other mineral nutrients such as Fe, Mg and Ca. stress reactions. The final step of abscisic acid 1676
1631 The critical toxicity level varies from 200 in maize biosynthesis is catalysed by Mo-enzyme alde- 1677
1632 to 5,300 ppm in sunflower (Foy et al. 1988). hyde oxidase, while sulphite oxidase protects the 1678
plant against toxic levels of sulphite. A novel 1679
1633 20.4.2.6 Molybdenum Mo-enzyme, mitochondrial amidoxime reducing 1680
1634 Most soils contain Mo between 0.6 and 3.5 ppm. component (mARC), has been reported on the 1681
1635 It occurs in soil as molybdate oxyanion, MoO42−. envelope of mammalian mitochondria. Here it is 1682
1636 A fraction of soil Mo also occurs in organic form. associated with detoxification and catalyses the 1683
reduction of N-hydroxylated amidines acting 1684
1637 20.4.2.6.1  Uptake and Distribution jointly with cytochrome b5 and cytochrome b5 1685
1638 Plants absorb Mo as molybdate ion (MoO42−). reductase. The homologues of this new enzyme 1686
1639 However, the uptake may be reduced due to the were found among plants and eubacteria. 1687
1640 competition by other anions like SO42−, whereas
1641 ions such as PO42− are known to enhance the 20.4.2.6.3  S  ymptoms of Deficiency 1688
1642 uptake of Mo. The uptake of Mo in plant cells has and Excess Mo 1689
1643 been discovered recently (Tejada-Jimenez et al. In Mo-deficient plants, particularly legumes, 1690
1644 2007; Tomatsu et al. 2007; Baxter et al. 2008). symptoms of N deficiency are common, produc- 1691
1645 Mo is cotransported through various mechanisms ing chlorosis in younger leaves and stunted 1692
1646 such as phosphate uptake system (Heuwinkel growth. In some dicotyledonous species, e.g. 1693
1647 et al. 1992), P-type ATPases, heavy metal trans- cauliflower, there is a drastic reduction in size 1694
1648 porters (Palmgren and Harper 1999), sulphate and irregularities in the formation of leaf blade. 1695
1649 transporters (Tweedi and Segel 1970) and non- Under such severe Mo deficiency, only the mid- 1696
1650 specific anion transporters (Mendel and Hansch rib of leaf is present giving it a whip appearance 1697
1651 2002). The chemical properties indicate that it is called ‘whiptail’. Local chlorosis and necrosis 1698
1652 transported as molybdate ion (MoO42−), similar to along the main veins of mature leaves called ‘yel- 1699
1653 sulphate (SO42−), phosphate (PO42−), tungstate low spots’ are common in citrus. 1700
1654 (WO42−) and vanadate (VO42−). A putative sul- Mo toxicity generally does not occur because 1701
1655 phate transporter, AtSultr5;2, has been identified there is a wide gap between the critical concen- 1702
1656 as molybdate transporter (MOT1) in Arabidopsis trations for deficiency and toxicity levels, which 1703
1657 (Tomatsu et al. 2007). MOT1 is classified as vary by a factor of 104 times (0.1–1,000 μg Mo 1704
1658 high-affinity transporter (Km <21 nM) expressed per gram dry weight). However, if at all it is taken 1705
1659 in both roots and shoots and located in plasma up beyond the toxic levels, malformation of 1706
1660 membrane and vesicles. In plants, Mo is localised leaves and a golden-yellow discoloration of shoot 1707
1661 mainly in the phloem and vascular parenchyma tissue occur. 1708
1662 and is readily translocated throughout the system
1663 (Gupta 1997). Unlike other elements, Mo can be 20.4.2.7 Chlorine 1709
1664 taken up in excess by the plants without resulting In nature, chlorine is abundantly present since it 1710
1665 in any toxic effects. is available in various sources such as soil 1711
 R. Pandey

1712 reserves, irrigation water, rain, fertilisers and air Mimosa pudica is directly controlled by this ion. 1759
1713 pollution. Therefore, in crop production, chloride The ‘osmotic motor’ for the leaf movement is 1760
1714 toxicity is of major concern than its deficiency. It powered by a plasma membrane H+-ATPase 1761
1715 is present in aqueous form in soil as monovalent which drives KCl and water fluxes. 1762
1716 anion, Cl−. It is not adsorbed to soil particles and
1717 is mobile thus highly susceptible to leaching loss. 20.4.2.7.3  S  ymptoms of Deficiency 1763
and Excess Cl− 1764
1718 20.4.2.7.1  Uptake and Distribution Cl deficiency symptoms include reduction in
−
1765
1719 Chlorine uptake is an active process and uptake leaf surface area, wilting of leaf at margins, inter- 1766
1720 occurs very rapidly in considerable amounts. veinal chlorosis of mature leaves and restricted 1767
1721 However, its uptake also takes place against an and highly branched root systems. Under severe 1768
1722 electrochemical gradient by carrier proteins pres- Cl− deficiency, curling of the youngest leaf fol- 1769
1723 ent in plasma membrane. In green tissue, under lowed by shrivelling and necrosis takes place. In 1770
1724 light Cl− uptake is enhanced as ATP formation palm trees, besides wilting and premature senes- 1771
1725 during photosynthetic phosphorylation provides cence of leaves, frond fracture and stem cracking 1772
1726 energy source for active uptake. Uptake of chlo- are typical Cl− deficiency symptoms. 1773
1727 ride through transporters was confirmed after the Toxicity of Cl− is a more serious problem in 1774
1728 discovery of chloride channel (CLC) family pro- agriculture. Cultivation on salt-affected soils 1775
1729 tein in A. thaliana and Nicotiana tabacum shows Cl− toxicity symptoms in plants. Typical 1776
1730 (Hechenberger et al. 1996, Lurin et  al. 1996). symptoms of Cl− toxicity are bronzing of leaf tips 1777
1731 Seven CLC genes each have been identified in A. and margins, premature yellowing and abscission 1778
1732 thaliana (AtClCa–AtClCg; Marmagne et  al. of leaves. Some crops such as sugar beet, barley, 1779
1733 2007) and rice (Oryza sativa, OsClC1–OsClC7; maize, spinach and tomato are highly tolerant, 1780
1734 Diédhiou and Golldack 2005). These CLC pro- while tobacco, beans, citrus, potatoes and 1781
1735 teins are involved in anion transport across plant legumes are highly susceptible to Cl− toxicity. 1782
1736 membranes (reviewed by De Angeli et al. 2009).
1737 Uptake of Cl− faces competition with NO3− 20.4.2.8 Nickel 1783
1738 and SO42−. Besides root, foliar absorption of Cl− Chemically Ni is related to Fe and cobalt. It 1784
1739 as chlorine gas also occurs. Chloride content in occurs not only in Ni(II) oxidation states but also 1785
1740 plant tissue is usually in the range of 50–500 μmol in I and III states in biological systems and forms 1786
1741 per kilogram of dry weight. In the cell, it is pref- stable complexes with cysteine, citrate and Ni 1787
1742 erentially stored in the vacuole. enzymes. Ni forms chelates and can readily 1788
replace other heavy metals (like Fe) from physi- 1789
1743 20.4.2.7.2  Biological Functions of Cl− ologically important sites. Ni content of soil is 1790
1744 Cl− is known to be associated with more than 130 usually less than 100 ppm. It is derived from the 1791
1745 organic compounds in plants. Cl− being a mobile weathering of ultrabasic igneous rocks (serpen- 1792
1746 anion, majority of its function is associated with tine mineral). 1793
1747 electrical charge balance. It plays an important
1748 role in photosynthesis. In photosystem II (P680) of 20.4.2.8.1  Biological Functions of Ni 1794
1749 oxygen evolving complex, there are three extrin- Nickel was the 17th mineral element to be 1795
1750 sic polypeptides which contain Mn. Cl− acts as a regarded as essential trace (micronutrient) ele- 1796
1751 bridging ligand and stabilises the oxidised state ment for plant growth. It was found to be a metal 1797
1752 of Mn in these associated polypeptides. H+- component in enzymes such as urease, dehydro- 1798
1753 pumping ATPase present on tonoplast is specifi- genases, hydrogenases and methyl reductases in 1799
1754 cally stimulated by Cl−. In guard cells of some a large number of bacteria. Earlier reports showed 1800
1755 plants, e.g. onion, Cl− accumulation regulates that low Ni concentration stimulates germination 1801
1756 opening and closing of stomata. Therefore, Cl− and growth of many crop species. The first evi- 1802
1757 indirectly affects plant growth by stomatal regu- dence of Ni as a constituent of urease enzyme in 1803
1758 lation. The seismonastic leaf movement of higher plants (jack bean) was provided by Dixon 1804
 20  Mineral Nutrition of Plants

1805 et al. (1975). Later, Ni requirement for legumes fusion of ions through the soil solution, (2) the 1847
1806 and nonlegumes was also established (Freyermuth passively charged ions carried along as water 1848
1807 et al. 2000). Some functions of Ni are now clearly moves by bulk flow into the roots and (3) extend- 1849
1808 defined and therefore it was included in the list of ing growth of roots towards the dissolved ions. In 1850
1809 essential micronutrients (Eskew et  al. 1983, both higher and lower plants, ion uptake is char- 1851
1810 1984). acterised by three principles: 1852
1811 Nickel is taken up as divalent cation, Ni2+. It is 1. Selectivity: Plants preferentially take up cer- 1853
1812 mobile in xylem and phloem, so after uptake, tain mineral elements, while others are dis- 1854
1813 considerable amount of it is transferred to the criminated against or almost excluded. In soil 1855
1814 seeds and fruits. Ni has an important role in N solution, some mineral elements may be pres- 1856
1815 metabolism. The urease enzyme present in plants ent in higher concentration, but they are not 1857
1816 is involved in breakdown of urea to CO2 and entirely required for plant growth. So, the 1858
1817 ammonia via the ornithine cycle. Accumulation plant has mechanism to exclude or selectively 1859
1818 of urea in the absence of urease would be toxic to take up only those elements required for its 1860
1819 the plant’s cellular machinery. Moreover, recent growth. 1861
1820 findings suggest that in addition to urea metabo- 2. Accumulation: The concentration of mineral 1862
1821 lism, plant ureases have a protective role against elements can be much higher in the plant cell 1863
1822 phytopathogens (Follmer 2008). sap than in the external solution. It has been 1864
seen that the ion concentration in the root cell 1865
1823 20.4.2.8.2  S  ymptoms of Deficiency sap is generally much higher than that in the 1866
1824 and Excess Ni nutrient solution. This is evident in the case of 1867
1825 In crop plants, Ni toxicity is a major problem potassium, nitrate and phosphate, which are 1868
1826 rather than deficiency. Under Ni deficiency, accu- accumulated at higher concentration in the 1869
1827 mulation of toxic levels of urea might occur. root cells. 1870
1828 However, the Ni toxicity symptoms closely 3. Genotype: Distinct differences among the 1871
1829 related to Fe deficiency symptoms. Acute Ni tox- plant species exist in terms of ion uptake. This 1872
1830 icity gives rise to chlorosis. In cereals, pale yel- difference depends on the condition of the 1873
1831 low stripes along the length of the leaf occur. The growing media. For example, in alga species, 1874
1832 whole leaf may turn white and necrosis occurs at Nitella grown in pond water had higher con- 1875
1833 leaf margins. In dicots, the Ni toxicity causes centration of potassium, sodium, calcium and 1876
1834 chlorotic patches between leaf veins, similar to chloride in the cell sap, while in Valonia 1877
1835 Mn deficiency. grown in highly saline sea water, only potas- 1878
sium remained at higher concentration in the 1879
cell sap, whereas the sodium and calcium con- 1880
1836 20.4.3 Beneficial Elements centrations remain at a lower level than in sea 1881
water. 1882
1837 Summary of beneficial elements regarding plant The accumulation of ions in plant tissues in 1883
1838 concentrations, beneficial effect, physiological quantities more than the circumambient solution 1884
1839 mechanisms and deficiency or toxicity symptoms indicates that the ions diffuse through the cell 1885
1840 of nutrients is presented in Table 20.6. against a concentration gradient. This type of ion 1886
uptake requires expenditure of metabolic energy, 1887
and therefore, it is dependent on the metabolic 1888
1841 20.5 Ion Absorption status of absorbing cells or tissues in plants. This 1889
phenomenon of ion or salt absorption, called 1890
1842 The anions and cations freely dissolved in the active absorption, and accumulation is most com- 1891
1843 soil solution are the most readily available forms mon in rapidly growing tissues such as meriste- 1892
1844 to be absorbed by the roots, even though per se matic cells, and it decreases with the increase in 1893
1845 their concentration may be low. These nutrients maturity of cells. There are some evidences 1894
1846 reach the vicinity of roots in three ways: (1) dif- which prove that the active absorption of solutes 1895
t6.1 Table 20.6  Plant concentrations, beneficial effect, physiological mechanisms and deficiency or toxicity symptoms of nutrients categorised as beneficial elements
t6.2 Element Form Plant concentration Beneficial effects Mechanisms hypothesised Symptoms Plant species/hyperaccumulators
t6.3 Aluminium Al3+, <0.1 % Increases root and shoot Increases antioxidant activityToxicity: inhibition of Miscanthus sinensis (maiden
t6.4 Al(OH)2+ non-accumulators growth Increases P availability root growth, alteration grass), Camellia sinensis (tea,
t6.5 Al(OH)2+ ≥0.1 % Al Resistance to herbivore Decreases Fe toxicity in root architecture, Melastoma malabathricum
t6.6 accumulators disruption of root
t6.7 elongation, symptoms
t6.8 similar to P deficiency
t6.9 Cobalt Co2+ 10−6 − 0.001 % Increases growth, nodule Cobalamin (vit B12)-dependent Deficiency: affects Families of Lamiaceae,
t6.10 Co3+ non-accumulators number and weight, enzyme systems in Rhizobium: nodule development, Scrophulariaceae, Asteraceae
t6.11 >0.1 % Co plant nutrient levels, as methionine synthase, yellowing of leaves and Fabaceae examples: Pisum
t6.12 accumulators well as seedpod yield ribonucleotide reductase, similar to N deficiency sativum (Pea), Lotus japonicus
t6.13 and seed quality methylmalonyl-coenzyme A symptoms, lower plant (Lotus) Crotalaria cobalticola
t6.14 in legumes mutase, inhibition of ethylene N content, (indicator plant)
t6.15 Retardation of leaf biosynthesis accumulation of Co in
t6.16 senescence root nodules
t6.17 Enhancement of drought Toxicity: symptom
t6.18 resistance in seeds resembles Mn
t6.19 Resistance to herbivore deficiency
t6.20 Sodium Na+ <0.05 % Increases plant growth, Cell expansion Enzymes activated by Atriplex vesicaria, Zea mays
t6.21 non-halophytes leaf area, number of Stomatal regulation by K are inhibited, less (maize), Sorghum bicolor
t6.22 >0.25 % in halophytes stomata per unit substituting K and maintaining starch content in tissue, (sorghum), members of
t6.23 leaf area higher relative water content accumulation of Chenopodiaceae
t6.24 Essential for C4 Regeneration of sucrose
t6.25 and CAM plants phosphoenolpyruvate
t6.26 Can facilitate
t6.27 nitrate uptake
t6.28 Improves water balance
t6.29 of plants
R. Pandey
t6.30 Selenium Se2−, SeO32−, ≤0.01 % non –Se Increases plant growth Increases antioxidant activity Toxic levels in plants Astragalus bisulcatus
t6.31 SeO42− accumulators Resistance to pathogen Prevents P toxicity lead to disorders in (milkvetch), species of
t6.32 0.01– 0.1 % Se and herbivore attack Replaces SO42− and forms animals feeding on Xylorrhiza, Stanleyea, Brassica
t6.33 accumulators Structural component of amino acids selenocysteine them; acute toxicity oleracea (broccoli) Ryegrass,
t6.34 ≥0.1 % Se specific selenoproteins (SeCys) and selenomethionine leading to death, lettuce, potato, duckweed
t6.35 hyperaccumulators and seleno-tRNAs (SeMet) chronic blindness and
t6.36 Volatile Se (dimethylselenide) paralyses, chronic
t6.37 deters herbivores, alkali disease leading
t6.38 Accumulated Se is toxic to to lameness
t6.39 herbivores and pathogens Deficiency of Se in
t6.40 produces symptoms of
20  Mineral Nutrition of Plants

t6.41 P toxicity in plants

t6.42 Silicon Si(OH)4 <0.5 % most species Maintains plant Strengthens cell walls In animals, fed on high Equisetum arvense (horsetail
t6.43 10–15 % horsetails sturdiness thus Activation or synthesis of Si content feed leads to millet), Oryza sativa (paddy
t6.44 preventing lodging, leaf stress-related molecules abrasion of rumen rice), Saccharum officinarum
t6.45 erectness for better light Systemic acquired resistance, wall, secondary (sugarcane)
t6.46 interception SAR-like bioactivity deposition in kidney
t6.47 Resistance to pathogen Regulation of metal transport
t6.48 and herbivore attack (Mn and Fe)
t6.49 Resistance to abiotic Increases antioxidant activity
t6.50 stress (heavy metals, Sodium exclusion from roots
t6.51 salinity, drought, UV
t6.52 radiation, extreme
t6.53 temperature)
 R. Pandey

1896 requires metabolic energy: (1) higher rate of res- This transpirational pull is so strong that water 1941
1897 piration increases salt accumulation inside the and nutrients move in upward direction to several 1942
1898 cell, (2) respiratory inhibitors check the process feet in the tallest trees growing in nature. Stout 1943
1899 of salt uptake and (3) by decreasing oxygen con- and Hoagland had proved that mineral nutrients 1944
1900 tent in the medium, the salt absorption also absorbed by the roots are translocated through 1945
1901 decreases. These evidences suggest that absorp- the xylem vessel. 1946
1902 tion of salt is directly coupled with respiratory Nutrient uptake by roots is a well-regulated 1947
1903 rate and energy level in the plant body. process and is also specific and selective. 1948
Subsequent translocation of ions across cortex to 1949
reach the stele (xylem loading) is also determined 1950
1904 20.5.1 Pathway of Solute by several interrelated factors. The pathway of 1951
1905 from External Solution ion or solute movement across the root faces 1952
1906 into the Cells some special constraints because of the anatomy 1953
of roots. All plant cells are separated by cell 1954
1907 The water and dissolved ions in the external solu- walls, and ions can move through channels in the 1955
1908 tion move into the xylem cells of roots via three cell wall spaces without ever entering a living 1956
1909 possible pathways: (1) through the cell walls or cell. This continuum of cell wall, which allows 1957
1910 apoplast of epidermal and cortical cells; (2) extracellular ion movement, is called the free 1958
1911 through the cytoplasmic or symplast system, space or apoplast. Besides cell wall forming a 1959
1912 moving from cell to cell; and (3) from vacuole to continuous phase, the cytoplasms of neighbour- 1960
1913 vacuole of the living root cells where the cytosol ing cells are also connected to each other and 1961
1914 of each cell forms part of the pathway. In the fol- form a continuum, collectively termed as the 1962
1915 lowing paragraphs, this pathway is discussed in symplast. The cytoplasmic bridges interconnect- 1963
1916 detail. ing the neighbouring cytoplasm are called plas- 1964
1917 Mineral salts dissolved in water are absorbed modesmata. These are cylindrical pores of 1965
1918 by roots and taken up through the xylem vessel 20–60 nm in diameter. Each plasmodesmata (sin- 1966
1919 from where it is distributed to all parts of the gular) is lined with a plasma membrane and con- 1967
1920 plant body. The dissolved cations and anions tains a narrow tubule, called desmotubule which 1968
1921 along with water moves from cell to cell through is the continuation of endoplasmic reticulum. 1969
1922 spaces between cell wall by diffusion, called apo- The cells near the root tips where most nutrient 1970
1923 plastic pathway. On the other hand, ions entering absorption occurs has a high density of 1971
1924 cell wall of the epidermis move across the cell plasmodesmata. 1972
1925 wall of cortex, cytoplasm of endodermis and cell The anatomy of root cell and the pathway of 1973
1926 walls of pericycle and finally reach the stele. This solute or ion movement in root cells are depicted 1974
1927 pathway involving the living (cytoplasm) part of in Fig. 20.7. The apoplast forms a continuous 1975
1928 the cell is referred to as symplastic pathway. In phase from the root surface through the cortex. 1976
1929 this pathway, nutrient elements entering the cyto- There is a layer of specialised cells called endo- 1977
1930 plasm of the epidermis move across the cyto- dermis at the boundary between vascular cylinder 1978
1931 plasm of the cortex and endodermis of pericycle and the cortex. The endodermis contains a layer 1979
1932 through plasmodesmata and finally reach the or strip of suberised cells known as Casparian 1980
1933 xylem vessels. strip, which functions as a barrier to the entry of 1981
1934 After the nutrients are absorbed by roots, it is water and mineral ions into the stele through the 1982
1935 translocated to other plant parts by the transpira- apoplast. This Casparian strip has hydrophobic 1983
1936 tion stream moving through the xylem. As water properties and completely surrounds each endo- 1984
1937 is continuously lost by aerial parts of the plant dermis cell. When a solute enters the root, it may 1985
1938 called transpiration, it creates a transpirational enter the symplast immediately by crossing an 1986
1939 pull. This driving force keeps the water moving epidermal cell plasma membrane, or it may dif- 1987
1940 up along with mineral salts in the xylem vessel. fuse between the epidermal cells through the cell 1988
 20  Mineral Nutrition of Plants

Fig. 20.7  Pathway of ion

transport across the root
Cortex
(A) symplastic
(B) apoplastic Stele
A
Early metaxylem

Late metaxylem
Root Phloem
hairs Endodermis
Casparian band

Exodermis
B (hypodermis)
Rhizodermis

1989 walls. From the apoplast of cortex, ions/solutes gradient and then leak out of the living cells of 2020
1990 may cross the plasma membrane of cortical cell, stele (possibly because of lower oxygen avail- 2021
1991 thus taking the symplastic route, or may radially ability interior of root) into the nonliving con- 2022
1992 diffuse all the way to endodermis through the ducting cells of xylem. 2023
1993 apoplastic route. In either case, the ion/solute The electrochemical potential of various ions 2024
1994 must first enter the symplast before entering into across the roots could be measured by using ion-­ 2025
1995 the stele due to the presence of Casparian strip. specific microelectrodes. From various studies, it 2026
1996 Once ions enter the symplast of root at epider- was indicated that K+, Cl−, Na+, SO42− and NO3− 2027
1997 mis or cortex, they are loaded into the tracheid or were all taken up actively by epidermal and corti- 2028
1998 vessel element of stele to be translocated to the cal cells and maintained in xylem against an 2029
1999 shoot. Since xylem treachery elements are dead electrochemical potential gradient when com- 2030
2000 cells, the ions exit symplast by crossing a plasma pared with the external medium. However, none 2031
2001 membrane for the second time. ‘Xylem loading’, of these ions is at a higher electrochemical poten- 2032
2002 as the name suggests, is the movement of ions tial in xylem than in cortex or living portion of 2033
2003 from symplast into the conducting cells of xylem. stele. Therefore, the final movement of ions into 2034
2004 Here again, the Casparian strip prevents back-­ xylem could be due to passive diffusion. However, 2035
2005 diffusion of ions through apoplast. Though this other studies have shown that this final step of 2036
2006 strip acts as a barrier to water and solute move- xylem loading may also involve active process 2037
2007 ment, the major advantage is that it helps the within the stele. By using treatments with inhibi- 2038
2008 plant to maintain an ionic concentration in xylem tors and other plant hormones, investigators have 2039
2009 higher than the soil water surrounding roots. shown that ion uptake by cortex and ion loading 2040
2010 There is a possibility that ions could enter tra- in xylem operates independently. Treatments 2041
2011 cheid and vessel element of xylem by passive dif- with protein synthesis inhibitor like cyclohexi- 2042
2012 fusion. But in this case, the movement of ions mide or with cytokinin (benzyladenine) inhibit 2043
2013 from root surface to the xylem would take only a xylem loading without affecting uptake by cor- 2044
2014 single step requiring metabolic energy. This sin- tex. This result indicated that efflux from the ste- 2045
2015 gle energy-dependent step of ion uptake occurs at lar cells is regulated independently from uptake 2046
2016 the plasma membrane surfaces of root epidermal, by cortical cells. 2047
2017 cortical and endodermal cells. According to the Recent biochemical studies have indicated 2048
2018 passive diffusion model, ions move passively into that the xylem parenchyma cells have a role in 2049
2019 the stele via symplast through a concentration xylem loading. The plasma membranes of xylem 2050
 R. Pandey

2051 parenchyma cells contain H+ pumps, water chan- space enables cortex cells to take up solutes 2092
2052 nels and a variety of ion channels specialised for directly from the external solution. 2093
2053 influx or efflux. Absorption of ions by roots is The cell walls contain negative charge because 2094
2054 pronounced in the root hair as compared to meri- of polygalacturonic acid [carboxylic groups (R. 2095
2055 stem or elongation zones. Presence of root hairs CHOO−)] present in the middle lamella. These 2096
2056 greatly increases the surface area available for negative charges in apoplasm act as cation 2097
2057 ion absorption. exchangers resulting in accumulation of cations, 2098
whereas the anions are repelled. Therefore, the 2099
entry of charged solutes is restricted in the free 2100
2058 20.5.2 Mechanisms of Ion space. Hope and Stevens (1952) introduced the 2101
2059 Absorption term apparent free space (AFS) to describe ‘free 2102
space’. AFS comprises of water free space (WFS) 2103
2060 According to Fick’s law, the movement of mole- which is freely accessible to ions, charged and 2104
2061 cules or ions by diffusion always takes place uncharged molecules. Another term, Donnan free 2105
2062 spontaneously from a place of high concentration space (DFS), was introduced where cation 2106
2063 to a low concentration, that is, down a concentra- exchange and anion repulsion take place 2107
2064 tion or chemical gradient until equilibrium is (Fig.  20.8). The various theories of passive 2108
2065 reached. This movement of molecules by diffu- absorption are discussed below: 2109
2066 sion without the involvement of metabolic energy
2067 is termed as passive transport. Once the equilib- 20.5.3.1 Mass Flow Hypothesis 2110
2068 rium is reached, no further net movement of sol- According to this theory, ions are absorbed by 2111
2069 ute will occur without the application of a driving root along with mass flow of water under the 2112
2070 force. On the other hand, movement of substances influence of transpiration. In detopped tomato 2113
2071 after the equilibrium is reached or against a con- plants, an increase in transpiration increased salt 2114
2072 centration gradient by involving metabolic absorption by removing ions after they were 2115
2073 energy termed as active transport. released into the xylem duct. The dilution thus 2116
caused enhanced ion absorption. This resulted in 2117
accumulation of a small proportion of total salt 2118
2074 20.5.3 Passive Absorption uptake by plants through passive absorption. 2119
However, this salt accumulation by passive pro- 2120
2075 The concept of passive absorption is based on cess could be explained as follows: (1) free diffu- 2121
2076 ‘outer space’ also called as ‘free space’ or ‘diffu- sion of ions along concentration gradient into the 2122
2077 sion space’. If only a portion of tissue volume is apparent free space of a tissue, (2) accumulation 2123
2078 open to free diffusion, the ions will move freely of ions against concentration gradient due to ion 2124
2079 in or out of the tissue. After sometime, the part of exchange or Donnan equilibrium and (3) mass 2125
2080 tissue undergoing free diffusion will reach equi- flow of ions through roots due to transpirational 2126
2081 librium with external solution resulting in same ‘pull’ may also occur. All these processes do not 2127
2082 ion concentration within the tissue as that of the require expenditure of metabolic energy. 2128
2083 external solution. The part of a cell or tissue that
2084 allows free diffusion of ions is called free space 20.5.3.2 Ion Exchange Theory 2129
2085 or outer space. Usually the volume of root tissue Both cations and anions have a tendency to get 2130
2086 accessible for free space is only a small fraction adsorbed on the surfaces of cell walls or mem- 2131
2087 ~5 % of total root volume. The extent of solute branes of cells and exchange with ions present in 2132
2088 flux into free space for a given volume of free soil solution. During the absorption of a posi- 2133
2089 space depends on several factors such as transpi- tively charged ion such as K+, either a positively 2134
2090 ration rate, solute concentration and root hair for- charged ion such as H+ is displaced from the cell 2135
2091 mation. However, the presence of such a free (ion exchange) or a negatively charged ion enters 2136
the cell. Similarly, anions can exchange with free 2137
 20  Mineral Nutrition of Plants

Fig. 20.8 Schematic -+ Indiffusible

representation of the pore
system of the apparent free -+ + - anions
space (Source: Marschner
1995)
- + +-
- - + -
+ + - -
- + +
- -
-
-

Water free
Macropore Donnan free space
space

Micropore
- Anion

+ Cation

2138 hydroxyl (OH−) ions. This process of exchange 3. Donnan equilibrium 2166
2139 between the adsorbed ions and ions in the solu- This theory takes into account the effect 2167
2140 tion is known as ion exchange. Sometimes the of fixed or nondiffusible ions. The cell con- 2168
2141 uptake of nutrient cations from a soil solution tents are separated from external solution by 2169
2142 into the roots exceeds the anion uptake and vice a differentially permeable membrane. This 2170
2143 versa. In each case, the neutrality is maintained. membrane is impermeable to anion concen- 2171
2144 In a case where anion uptake exceeds cation tration present in cell. A negative nondiffus- 2172
2145 uptake, the OH- and bicarbonate (HCO3−) ions ing charge on one side will create a potential 2173
2146 are transported outwardly from inside the cells gradient across the membrane through which 2174
2147 into the free space. Likewise, if cation absorption the ions will diffuse. This results in diffusion 2175
2148 exceeds anion uptake, cells exchange some of equal number of cation and anion through 2176
2149 hydrogen ions. The ionic exchange process is the membrane until electrochemical equilib- 2177
2150 explained as follows: rium is reached. However, because of the 2178
2151 1. Carbonic acid exchange theory ‘fixed’ negative (anion) charge on the inner 2179
2152 The roots respire continuously giving out side of the membrane, additional cations 2180
2153 CO2 in the rhizosphere. This combines with will be required to balance this charge. 2181
2154 water and forms carbonic acid (H2CO3), which Therefore, the cation concentration will be 2182
2155 dissociates into H+ and HCO3− ions. A zone of greater in internal solution (inside the cell) 2183
2156 carbonic acid is developed in root tips because than the external solution. Also, the concen- 2184
2157 of high respiratory activity. The H+ ions tration of anions in external solution will be 2185
2158 exchange with cations absorbed on the clay less than that of the external one because of 2186
2159 micelle, and the cations come into soil solu- the excess of negative charges due to ‘fixed’ 2187
2160 tion where it diffuses onto the root surface. anions. Therefore, Donnan equilibrium, a 2188
2161 2. Contact exchange theory term proposed by F.G. Donnan, is attained if 2189
2162 A similar ion exchange takes place between the product of anions and cations in the 2190
2163 root and soil colloids at the point where they internal solution becomes equal to the prod- 2191
2164 are in direct contact with each other without uct of anions and cations in the external 2192
2165 being first dissociated in the soil solution. solution. 2193

Donnan equilibrium =
( Positive ions inside ) = ( Negative ions inside )
( Positive ions outside ) ( Negative ions outside ) 2194
 R. Pandey

2195 20.5.3.3 Facilitated Diffusion be discussed in the following sections. Active 2212
2196 The passive transport can also take place by car- transport is further divided into two categories: 2213
2197 rier proteins localised in the plasma membrane primary active transport and secondary active 2214
2198 where it can transport a much wider range of pos- transport. 2215
2199 sible substances. This passive transport mediated
2200 by a carrier is called facilitated diffusion. 20.5.4.1 Primary Active Transport 2216
2201 However, it resembles diffusion only in that it When the transport of ions is directly coupled to 2217
2202 transports substances down their gradient of elec- expenditure of metabolic energy, such as ATP 2218
2203 trochemical potential without any additional hydrolysis, an oxidation-reduction reaction (the 2219
2204 input of energy. electron transport chain of mitochondria and 2220
chloroplast), or absorption of light by the carrier 2221
protein (in halobacteria, bacteriorhodopsin), it is 2222
2205 20.5.4 Active Absorption termed as primary active transport. Membrane 2223
proteins involved in primary active transport are 2224
2206 As discussed in the previous section, active called pumps. Most pumps transport energy ions, 2225
2207 absorption involves transport of solutes against such as H+ or Ca2+. 2226
2208 the concentration gradient. Carriers, pumps and The pumps can be further characterised as 2227
2209 channels located in the membrane are involved in either electrogenic or electroneutral. Electrogenic 2228
2210 active transport of ions/solutes (Fig. 20.9). It transport refers to ion transport involving the net 2229
2211 involves electrogenic and cytochrome pumps to movement of charge across the membrane, 2230

Plasma
membrane Tonoplast
Apoplasm Cytoplasm Vacuole

Uniport Cation
Cation
Counter
Cation transport
(K+)
Co-
Co-transport
H+
ATP
ATP
H+- H+-
+ ATPase + ATPase
H H H+ H+
(pump) (pump)
ADP + Pi
ADP + Pi
+
H
Co-
Co-transport
Anion Uniport
Anion
FeIII NAD(P)H
PPi
e-
H+ TM redox
pump PPiiase
H+ H+
H+ (pump)
FeII NAD(P)+ 2Pi

Ion channels Ion channels

pH ~5.5 pH 7.3 – 7.6 pH 4.5 – 5.9

Fig. 20.9  Model for the localisation and functioning channels and transport of cations and anions across the
of electrogenic proton pumps (H + -ATPase, PPiase), plasma membrane and tonoplast (Source: Marschner
transmembrane redox pump (NAD(P) oxidase), ion 1995)
 20  Mineral Nutrition of Plants

2231 whereas in electroneutral transport, no net move- created when protons are extruded from the 2278
2232 ment of charge is involved. The pumps are driven cytosol by the electrogenic H+-ATPases. This 2279
2233 by energy released during hydrolysis of ATP car- takes place both at the plasma membrane and at 2280
2234 ried out by ATPase located at plasma membrane the vacuolar membrane where a membrane 2281
2235 and tonoplast. This enzyme is considered as potential and pH gradient develop at the expense 2282
2236 ‘master enzyme’ because of its key role in regula- of ATP hydrolysis. The PMF represents stored 2283
2237 tion of cytoplasmic pH and the driving force for free energy or the H+ gradient that is used to drive 2284
2238 cation and anion uptake. For example, the Na+/ the transport of other substances against their 2285
2239 K+-ATPase of animal cell pumps out three Na+ chemical potential gradient. The secondary active 2286
2240 for every two K+ ions taken in, resulting in net transport may be either symport (two ions/solutes 2287
2241 outward movement with one positive charge. The moving in the same direction) or antiport (down- 2288
2242 Na+/K+-ATPase is therefore an electrogenic ion hill proton movement drives active (uphill) trans- 2289
2243 pump. In contrast, H+/K+-ATPase of the animal port of solute in the opposite direction). Transport 2290
2244 gastric mucosa pumps one H+ out of the cell for by carriers may be 100–1,000 ions or molecules 2291
2245 every one K+ taken in, so there is no net move- per second. 2292
2246 ment of charge across the membrane. Therefore, The presence of ion channels in plant cell 2293
2247 the H+/K+-ATPase is an electroneutral pump. membranes has also been established. These ion 2294
2248 In general, H+ is the principal ion which is channels have a unique ability to regulate or 2295
2249 electrogenically pumped across the cell mem- ‘gate’ ion flux subject to the physical and chemi- 2296
2250 branes of plants, fungi and bacteria. The plant cal environment of the channel protein. Transport 2297
2251 H+-ATPases exhibit a regulatory role in creating through the channel may or may not involve 2298
2252 the electrochemical potential gradient across the ­transient binding of solute to the channel protein. 2299
2253 plasma membrane, while the vacuolar H+-ATPase In any case, as long as the channel pore is open, 2300
2254 (V-ATPase) and the H+ pyrophosphatase (H+- solute that can penetrate the pore diffuses through 2301
2255 ppase) pump proton into the lumen of vacuole it extremely rapidly. Ion transport through chan- 2302
2256 and Golgi cisternae. nels is always passive. The pore size and density 2303
of charges on interior lining determine the trans- 2304
2257 20.5.4.2 Secondary Active Transport port specificity; channel transport may be mainly 2305
2258 The other important mechanism by which solutes limited to ions or water. The region in the channel 2306
2259 can be transported across membrane against the that determines specificity is called ‘selectivity 2307
2260 gradient of electrochemical potential is coupling filter’. However, ion channels remain closed most 2308
2261 of ‘uphill’ transport of one ion/solute to the of the time and their number per cell is also 2309
2262 ‘downhill’ transport of another. Carrier proteins meagre. 2310
2263 located in the membranes carry out such cotrans- A class of proteins that form water channels, 2311
2264 port of ions/solutes, which is also called ‘second- relatively abundant in the membranes, are named 2312
2265 ary active transport’. These carriers are indirectly ‘aquaporins’. Aquaporins are common to plant 2313
2266 driven by the action of pumps. In carrier-­mediated and animal membranes; their expression and 2314
2267 ion transport, the substance being transported is activity in response to water availability are regu- 2315
2268 initially bound to a specific site on the carrier lated by several mechanisms including protein 2316
2269 protein. This binding causes a conformational phosphorylation. 2317
2270 change in the protein, which exposes the sub-
2271 stance to solution on the other side of the mem- 20.5.4.3 Cytochrome Pump 2318
2272 brane. Transport completes when substance H.  Lundegardh in 1954 proposed this theory 2319
2273 dissociates from the carrier’s binding site. Rate of when he observed a quantitative relationship 2320
2274 transport by a carrier is about 106 times slower between anion absorption and respiration, 2321
2275 than transport through a channel. whereas no such correlation with cation absorp- 2322
2276 Secondary active transport utilises the energy tion was found. It was also noticed that salt respi- 2323
2277 stored in the proton-motive force (PMF) or ∆p, ration and anion absorption were inhibited by 2324
 R. Pandey

2325 cyanide or even carbon monoxide. He, therefore, intermediates. The rate of respiration solely 2330
2326 suggested that anions could be transported across determined by anion absorption is called ‘anion 2331
2327 the membrane by cytochrome system and absorp- respiration’ or ‘salt respiration’. The respiration 2332
2328 tion of anion is independent of cation. Energy is sup- rate (excluding anion respiration) observed in 2333
2329 plied by direct oxidation of respiratory distilled water is called ‘ground respiration’. 2334

2335
( ) ( )
Total respiration Rt = Ground respiration Rg + Salt or anion respiration ( Ra )

2336 20.5.5 Passive Absorption

are abundant in the natural environment. Use of 2371
biofertilisers leads to enhanced crop productivity 2372
2337 The absorption of minerals without expenditure and reduction in the usage of chemical fertilisers 2373
2338 of metabolic energy is termed ‘passive absorp- resulting in improvement of soil texture and miti- 2374
2339 tion’. Passive ion absorption in the root system gating other harmful environmental effects. As a 2375
2340 was demonstrated by Briggs and Robertson matter of fact, the system of integrated nutrient 2376
2341 (1957). The distinguishing characteristics of pas- management (INM) subscribes to the use of a 2377
2342 sive absorption include: combination of inorganic and organic/biofertilis- 2378
2343 1. Mineral salt absorption not being affected by ers. Biofertilisers include the following groups: 2379
2344 temperature or metabolic inhibitors 1. Symbiotic N fixers, e.g. Rhizobium spp. for 2380
2345 2. Rapid uptake of ions when plants are trans- legumes 2381
2346 ferred from a medium of low to high 2. Non-symbiotic free-living N fixers, e.g. 2382
2347 concentration Azotobacter, Azospirillum 2383
3. Algae biofertilisers (blue-green algae (BGA) 2384
in association with Azolla) 2385
2348 20.6 Biofertilisers 4. Phosphate solubilising bacteria (PSBs) 2386
5. Mycorrhiza 2387
2349 Biofertilisers are natural fertilisers consisting of 6. Organic fertilisers 2388
2350 microbial inoculants of bacteria, fungi and algae,
2351 either alone or in combination resulting in
2352 enhanced growth of plants mediated by increased 20.6.1 Symbiotic Nitrogen Fixers 2389
2353 availability of nutrients. Biofertilisers can be and Algal Biofertilisers 2390
2354 applied to seed, plant surfaces or soil, where they
2355 colonise the rhizosphere or the interior of plant Bacteria such as Rhizobium spp. exhibit symbi- 2391
2356 and increase the supply of primary nutrients to otic relationship with host plants especially 2392
2357 the host plant. These microorganisms, either legumes. Blue-green algae (BGA) or cyanobac- 2393
2358 through symbiotic or non-symbiotic association, teria have been used in agriculture since long. 2394
2359 help the plants to fix N from atmosphere, solu- The bacterial biofertilisers fix about 50–150  kg 2395
2360 bilise and mobilise fixed P; translocate minor ele- N/ha or more per year. The root-nodulating bac- 2396
2361 ments like Zn and Cu to the plants; produce plant teria, such as Rhizobium, are common in nature. 2397
2362 growth-promoting hormones, vitamins and However, there are a few genera that produce 2398
2363 amino acids; and control plant pathogenic fungi. stem nodules. For example, Azorhizobium spp. 2399
2364 Thus, biofertilisers are expected to reduce the use produce stem nodules on Sesbania rostrata. 2400
2365 of chemical fertilisers and pesticides. Besides Rhizobium is classified based on the specific 2401
2366 microorganisms, biofertilisers include organic legume species they nodulate. For example, R. 2402
2367 fertilisers such as farmyard manure (FYM) leguminosarum nodulates pea (Pisum sativum), 2403
2368 obtained by decomposing farm wastes. khesari (Lathyrus sativus) and lentil (Lens culi- 2404
2369 The use of biofertilisers is both economical naris); R. phaseoli produces nodules on French 2405
2370 and environment friendly. These microorganisms bean (Phaseolus vulgaris) and bean (Phaseolus 2406
 20  Mineral Nutrition of Plants

2407 multiflorus); R. meliloti nodulates lucerne to the bacterial genera Thiobacillus, Bacillus, etc. 2449
2408 (Medicago sativa) and fenugreek (Trifolium These bacteria, known as phosphate solubilising 2450
2409 foenumgraecum); R. trifoli nodulates clover bacteria (PSBs), produce substances like pseudo- 2451
2410 (Trifolium sp); and R. lupini nodulates white bactin (also called ‘siderophores’) that 2452
2411 lupins (Lupinus alba). chelate Fe. 2453
2412 Cyanobacteria or BGA are photosynthetic Availability of P is low both in alkaline and 2454
2413 prokaryotic organisms that can fix atmospheric N acidic soils since it is fixed as insoluble oxides. 2455
2414 in association with a fresh water fern called The inoculants of PSBs when applied to either 2456
2415 Azolla. Cyanobacteria belong to the genera soil or seed can be useful to mobilise fixed 2457
2416 Nostoc, Anabaena, Plectonema, Aulosira and P. Rock phosphate when used with PSBs can 2458
2417 Tolypothrix. Azolla cultures are inoculated in reduce the phosphatic fertiliser requirement by 2459
2418 paddy fields under both aerobic and lowland con- 50 %. Mere seed inoculation with PSBs results in 2460
2419 ditions. However, the effectiveness of the inocu- crop yield responses equivalent to that obtained 2461
2420 lum is higher under lowland condition, wherein by application of 30 kg P2O5 per ha in the form of 2462
2421 about 20–30 kg N is fixed per ha per crop season. chemical fertilisers. 2463
2422 The enzyme nitrogenase is present in cyanobac-
2423 teria and N fixation occurs in specialised cells
2424 called ‘heterocysts’. These heterocysts have nif 20.6.4 Mycorrhizae 2464
2425 gene (involved in N fixation) and also act as
2426 oxygen-­proof compartment, protecting nitroge- Mycorrhizae (mycor fungus; rhiza root) are a 2465
2427 nase from oxygen inactivation. Azolla owes its symbiotic (intimate) and mutualistic (mutually 2466
2428 N-fixing capacity to Anabaena, while the fern beneficial) association between a nonpathogenic 2467
2429 supplies food material to the alga. In addition to or weakly pathogenic fungus and living root cells 2468
2430 N fixation, BGA decomposition contributes to of plant, particularly cortical and epidermal cells. 2469
2431 the ecosystem biomass, thereby enhancing soil The fungi receive organic nutrients from the plant 2470
2432 physical properties. and, in turn, improve the mineral element and 2471
water-absorbing properties of roots. The roots of 2472
most soil grown plants are mycorrhizal. About 2473
2433 20.6.2 Non-symbiotic Free-Living N 83 % of dicot and 79 % of monocot plants and all 2474
2434 Fixers gymnosperms are mycorrhizal worldwide. 2475
Generally, the fungus infects tender young roots 2476
2435 Azotobacter and Azospirillum are free-living bac- because, on older parts, the epidermis and cortex 2477
2436 terial genera which fix atmospheric N. are lost and a protective layer of suberin develops 2478
2437 Azotobacter can be used to inoculate crops like in cork cells. Root hair production either slows or 2479
2438 maize, wheat, mustard, cotton, potato and other ceases upon infection, so mycorrhizae often have 2480
2439 vegetable crops. By utilising soil organic matter, few such hairs. This reduction in root hairs 2481
2440 it fixes about 30 kg N per ha per year. Azospirillum greatly decreases the absorbing surface area; 2482
2441 inoculants are recommended especially for culti- however, the slender fungal hyphae extending 2483
2442 vation of wheat, maize, sugarcane, millets and from mycorrhizae explore greater soil volume. 2484
2443 sorghum. Two main groups of mycorrhiza are present in 2485
nature: (1) ectomycorrhiza and (2) endomycor- 2486
rhiza. However, there is a rare group with inter- 2487
2444 20.6.3 Phosphate Solubilising mediate properties called the ectendotrophic, 2488
2445 Microorganisms which is also sometimes found. In ectomycorrhi- 2489
zae (ECM), the fungal hyphae form a mantle out- 2490
2446 Phosphate solubilising microorganisms capable side the root and also inside the root in the 2491
2447 of solubilising Pi from insoluble sources enhance intercellular spaces of epidermis and cortex. No 2492
2448 P bioavailability. These microorganisms belong intracellular penetration into epidermal or corti- 2493
 R. Pandey

2494 cal cells occurs, but an extensive network called (increase drought tolerance capacity) and 2541
2495 the Hartig net is formed between these cells. suppresses soil-borne fungal and bacterial root 2542
2496 Ectomycorrhizae are commonly found on the pathogens (e.g. Pseudomonas syringae in tomato 2543
2497 roots of trees including members of Pinaceae plant). 2544
2498 family (pine, fir, spruce, larch, hemlock),
2499 Fagaceae (oak, beech, chestnut), Betulaceae
2500 (Birch, alder), Salicaceae (willow, poplar) and a 20.6.5 Organic Fertilisers 2545
2501 few other families.
2502 In endomycorrhiza, fungi live inside the corti- Organic fertilisers are the plant and animal wastes 2546
2503 cal cells and also grow in the intercellular spaces. which after decomposition release nutrients 2547
2504 Endomycorrhiza consists of three subgroups, essential for plant growth. These include farm- 2548
2505 namely, the vesicular arbuscular mycorrhizae yard manures (decomposed mixture of dung, 2549
2506 (VAM) the most common, the ericoid and the urine and litter of farm animals), compost (rotted 2550
2507 orchidaceous mycorrhizae. The VAM belongs to wastes of farm like sugarcane trash, paddy straw, 2551
2508 the family Endogonacae. They produce a etc.), sewage and sludge, vermicompost (decom- 2552
2509 branched haustorial structures called arbuscules, position of organic matter by earthworms), green 2553
2510 between the cortical cells and a mycelium that manures (undecomposed plant material) and 2554
2511 extends out in the soil. These extraradical myce- other livestock manures (poultry, sheep and goat 2555
2512 lium or hyphae are involved in absorbing mineral sweepings). These organic manures contain 2556
2513 salts and water. The arbuscules are short lived, small percentage of nutrient elements and they 2557
2514 about 10–12 days, and are the main sites of solute are applied in large quantities. The practice of 2558
2515 exchange with the host. There are mainly four growing crops with only organic fertilisers and 2559
2516 genera of VAM fungi, namely, Acaulospora, without the use of chemical pesticide or ­herbicide 2560
2517 Gigaspora, Glomus and Sclerocystis. Out of is called organic farming. Besides supplying 2561
2518 these genera, Glomus is present abundantly in nutrients, the organic fertilisers improve soil 2562
2519 soil. It has been found that not all endomycor- physical properties, increase availability of other 2563
2520 rhizal fungi produce vesicles as lipid-rich storage nutrients and control plant parasitic nematodes 2564
2521 organs, so the VAM has been renamed as AM and fungi. 2565
2522 (arbuscular mycorrhiza).
2523 The host plants secrete root exudates contain-
2524 ing flavanoids (e.g. β-estradiol), which attract 20.6.6 Advantages and Limitation 2566
2525 these fungi, and the root infection and colonisa- of Biofertilisers 2567
2526 tion occur. In mycorrhizal roots, for fungal
2527 growth, a substantial proportion of carbons fixed The relevance of biofertilisers is increasing rap- 2568
2528 during photosynthesis are required. In AM plants, idly. This is because it has been realised that 2569
2529 root respiration may be 20–30 % higher than in application of chemical fertilisers causes serious 2570
2530 non-mycorrhizal plants, and 87 % of higher res- harmful effects on the environment and human 2571
2531 piration is attributed to the fungus. The mycor- health. Further, they are costly and in short sup- 2572
2532 rhizal colonisation affects root and shoot growth ply. Therefore, the advantages of using biofertil- 2573
2533 differently. Its effect is more pronounced in isers in general are as follows: 2574
2534 nutrient-­poor soils rather than nutrient-rich soils. 1. They add nutrients to the soil and/or increase 2575
2535 Mycorrhizal colonisation helps the plant to their availability to the crops. 2576
2536 acquire nutrients which are less mobile in soil, 2. They secrete certain growth-promoting sub- 2577
2537 particularly P and also N, Zn, Cu and S. Besides stances such as indole acetic acid (IAA). 2578
2538 these, the mycorrhizal association also helps in 3. Under certain conditions, they exhibit antifun- 2579
2539 increasing the hormonal (IAA, cytokinin, ABA) gal activities, thereby protecting the plants 2580
2540 content of plants, improves plant-water relations from pathogenic fungi. 2581
 20  Mineral Nutrition of Plants

2582 4. They are harmless and eco-friendly low-cost other plant nutritionists, Dennis R. Hoagland and 2627
2583 agro-input supplementary to chemical Daniel I. Arnon, at the University of California, 2628
2584 fertilisers. wrote a classic in 1938  in agricultural bulletin 2629
2585 5. They improve soil structure (porosity) and ‘The Water Culture Method for Growing Plants 2630
2586 water-holding capacity. without Soil’. They developed several formula- 2631
2587 6. They enhance seed germination. tions for mineral nutrient solutions, known as 2632
2588 7. They increase soil fertility and fertiliser use Hoagland solutions and the modified Hoagland 2633
2589 efficiency and ultimately increase yield by solutions that are still in use today. This tech- 2634
2590 15–20 % in general. nique was used to establish the criteria of essenti- 2635
2591 However, the limitation in the use of biofertil- ality for different nutrient elements. 2636
2592 isers lies in the fact that its acceptability by the There are primarily two types of hydroponics, 2637
2593 farming community is low, the reason being viz., (1) solution culture and (2) medium culture. 2638
2594 organic fertilisers do not produce quick and spec- In solution culture, there is no solid medium for 2639
2595 tacular responses. Moreover, the quantity of the roots to support, just the nutrient solution, 2640
2596 nutrients provided by biofertilisers is not suffi- while the medium culture has a solid medium for 2641
2597 cient to adequately meet the crop demand for the roots and is named after the type of medium 2642
2598 high yields. being used. Again, solution culture is divided 2643
into three main methods of growing plants: 2644
(a) Static solution culture – plants are grown in [AU6]
2645
2599 20.7 Hydroponics: History containers of nutrient solution, such as plas- 2646
2600 and Scope in Indian tic buckets, tubs or tanks. The solution may 2647
2601 Agriculture or may not be aerated, but usually gentle 2648
aeration is required. 2649
2602 Hydroponics (Greek words ‘hydro’ water; (b) Continuous flow solution culture – the nutri- 2650
2603 ‘ponos’ labour) is a method of growing plants ent solution constantly flows past the roots, 2651
2604 using mineral nutrient solutions without soil. It is e.g. nutrient film technique. 2652
2605 also called “controlled environment agriculture” (c) Aeroponics  – no substrate is required, and 2653
2606 (CEA) since raising plants hydroponically the roots are suspended in the air in a closed 2654
2607 requires control of environmental factors such as chamber and are sprayed intermittently with 2655
2608 light intensity and duration, temperature, humid- fine drops of mist or aerosol of nutrient solu- 2656
2609 ity, pH of the solution/medium and mineral tion. Aeroponics is widely used in laboratory 2657
2610 nutrients. studies of plant physiology. National 2658
2611 The study of crop nutrition began thousands Aeronautics and Space Administration 2659
2612 of years ago. The classic work on growing ter- (NASA) has done extensive hydroponic 2660
2613 restrial plants without soil was published by Sir research for their ‘Controlled Ecological Life 2661
2614 Francis Bacon in 1627, the book named ‘Sylva Support System’ (CELSS). Special attention 2662
2615 Sylvarum’. After Bacon’s work, water culture has been given to this technique since a mist 2663
2616 became a popular research technique. John is easier to handle in a zero-gravity environ- 2664
2617 Woodward (1699) published his work on water ment as compared to a liquid. 2665
2618 culture experiments with spearmint where he In medium culture, there are variations for 2666
2619 mentioned that ‘plants grew better in less pure each media, i.e. passive subirrigation, flood and 2667
2620 water sources than plants in distilled water’. drain subirrigation, top irrigation and deep-water 2668
2621 German botanists, Julius von Sachs and Wilhelm culture. The media used for growing plants may 2669
2622 Knop (1859–1865), developed the techniques of be expanded clay, rock wool, coir, perlite, ver- 2670
2623 soilless cultivation. It was Professor William miculite, sand, gravel/quartz and brick shards. 2671
2624 Frederick Gericke (1937) who finally introduced Plant nutrients used in hydroponics are dis- 2672
2625 the term hydroponics and wrote the book named solved in water and are mostly in inorganic and 2673
2626 Complete Guide to Soilless Gardening. Two ionic forms. All the 17 elements that are essential 2674
 R. Pandey

2675 for plant growth are supplied using different Barber J (2003) Photosystem II: the engine of life. Q Rev 2721
Biophys 36:71–89 2722
2676 chemical combinations. Chelating agents such
Baxter I, Muthukumar B, Park HC, Buchner P, Lahner B 2723
2677 as EDTA are used to keep Fe in soluble form. et al (2008) Variation in molybdenum content across 2724
2678 The pH of nutrient solution ranges from 5.6 to broadly distributed populations of Arabidopsis thali- 2725
2679 6.0 depending on the crop. Once the plants are ana is controlled by a mitochondrial molybdenum 2726
transporter (mot1). PLoS Genet 4(2):e1000004 2727
2680 grown in nutrient solution, the composition of
Briat JF, Lobreaux S (1998) Iron storage and ferritin in 2728
2681 solution is altered due to depletion of specific plants. Met Ions Biol Syst 35:563–584 2729
2682 nutrients more rapidly than others, absorption of Briggs GE, Robertson RN (1957) Apparent free space. 2730
2683 water from the solution and alteration of pH by Annu Rev Plant Physiol Plant Mol Biol 8:11–30 2731
Buchner P, Takahashi H, Hawkesford MJ (2004) Plant 2732
2684 excretion of either acidity or alkalinity. Utmost
sulphate transporters: co-ordination of uptake, intra- 2733
2685 care should be taken while deciding the optimum cellular and long-distance transport. J Exp Bot 2734
2686 salt concentration to be used in solution, so as to 55(404):1765–1773 2735
2687 avoid toxicity, early nutrient depletion or pH Curie C, Cassin G, Couch D, Divol F, Higuchi K et  al 2736
(2009) Metal movement within the plant: contribution 2737
2688 changes which may together attribute to inadver-
of nicotianamine and yellow stripe 1-like transporters. 2738
2689 tent effects on plant growth and development. Ann Bot 103:1–11 2739
2690 Hydroponics was not a popular practice in De Angeli A, Monachello D, Ephritikhine G, Frachisse J-M, 2740
2691 India until 1946. The Government of Bengal ini- Thomine S et al (2009) CLC-mediated anion transport in 2741
plant cells. Philos Trans R Soc B 364:195–201 2742
2692 tiated Experimental Farm at Kalimpong,
Demidchik V, Maathuis FJM (2007) Physiological roles 2743
2693 Darjeeling. After a careful appraisal of salient of non-selective cation channels in plants: from salt 2744
2694 problems, the Bengal System of Hydroponics stress to signalling and development. New Phytol 2745
2695 was developed in 1946–1947 representing the 175:387–404 2746
Diédhiou CJ, Golldack D (2005) Salt-dependent regula- 2747
2696 efforts to meet Indian requirements. Recently, in
tion of chloride channel transcripts in rice. Plant Sci 2748
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