Iron distribution in vine leaves with HCO 3 induced chlorosis

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  • This article was downloaded by: [Moskow State Univ Bibliote]On: 26 August 2013, At: 16:41Publisher: Taylor & FrancisInforma Ltd Registered in England and Wales Registered Number:1072954 Registered office: Mortimer House, 37-41 Mortimer Street,London W1T 3JH, UK

    Journal of Plant NutritionPublication details, including instructions forauthors and subscription information:http://www.tandfonline.com/loi/lpla20

    Iron distribution in vineleaves with HCO3

    inducedchlorosisK. Mengel a , W. Bubl a & H. W. Scherer aa Institute of Plant Nutrition, Justus LiebigUniversity, Giessen, West GermanyPublished online: 21 Nov 2008.

    To cite this article: K. Mengel , W. Bubl & H. W. Scherer (1984) Iron distributionin vine leaves with HCO3

    induced chlorosis, Journal of Plant Nutrition, 7:1-5,715-724

    To link to this article: http://dx.doi.org/10.1080/01904168409363236

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  • JOURNAL OF PLANT NUTRITION, 7(1-5), 715-724 (1984)

    IRON DISTRIBUTION IN VINE LEAVESWITH HCO3- INDUCED CHLOROSIS

    KEY WORDS: Iron chlorosis, iron accumulation, bicarbonate, grapevine plants

    K. Mengel, W. Bubl and H. W. Scherer

    Institute of Plant NutritionJustus Liebig University

    Giessen, West Germany

    ABSTRACT

    The objective of the investigation was to examine whether ironchlorosis in grape vine grown on calcareous soils was related tothe Fe distribution in the leaf.

    Leaf samples collected from three different sites showed inmost cases higher Fe contents in the chlorotic leaves as comparedwith healthy leaves. The solubility of leaf Fe in diluted HC1, how-ever, was lower in chlorotic leaves than in green leaves.

    Enzymatic dissolution of leaves into vascular tissue, inter-costal cells and chloroplasts revealed that the Fe content in theintercostal cells of green leaves was significantly higher than inthe intercostal cells of chlorotic leaves. In addition, the inter-costal cells of chlorotic leaves had extremely high Ca and P con-tents.

    The P content of green and chlorotic leaves was not related tothe level of available P 1n the soil. It is, therefore, concludedthat the high P content in chlorotic leaves is the sequence and notthe cause of Fe chlorosis.

    715

    Copyright 1984 by Marcel Dekker, Inc. 0190-4167/84/0705-0715$3.50/0

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  • 716 MENGEL, BUBL, AND SCHERER

    On each of the three sites investigated, higher clay contentswere found under chlorotic grape vine plants than under healthyones. It is assumed that because of this higher clay content, soilcompaction may occur, resulting in an accumulation of C02 and in anincrease of the HCO3- concentration in the soil solution.

    INTRODUCTION

    According to recent investigations Fe chlorosis ts occurringvery often in connection with high concentration of HCO3" in thesoil (Boxma , Kovanci et al. 8). Although it has been shown that theuptake and translocation of iron in the plants is affected by HCO3"(Rutland and Bukovac^), to control chlorosis more information isrequired about the process which is leading to the HCO3" inducedchlorosis. Frequently total Fe in chlorotic leaves of plants grownon calcareous soils is higher than the Iron content in green leaves(Bubl3), in contrast to the HC1 soluble Fe which is lower in chlo-rotic leaves as compared with green leaves (Jacobson', Bucher^).

    The objective of the investigation presented here was to findout whether HCO3" induced chlorosis in grape vine was related tothe distribution of iron in the leaves. For this purpose chloroticand green leaves of vine plants were sampled and analyzed for dif-ferent Fe fractions: Total Fe, HC1 soluble Fe, Fe of the inter-costal cells (cells between leaf veins), Fe of the main leaf veins,and chloroplast Fe.

    MATERIALS AND METHODS

    In 1979 leaf samples from chlorotic and healthy vine plantswere collected from the site 'SpieBheim' and in 1980 from the sites'SpieBheim', 'Iphofen' and 'Nierstein1. These sites are susceptibleto Fe chlorosis of vine. On each site leaf samples were taken from20 green and 20 chlorotic vine plants, respectively. From each ofthese plants the 10 youngest leaves of two shoots were collected.The leaf samples were frozen immediately in liquid nitrogen and thenstored in a deep-freezer at -18C.

    Five samples were used for the determination of the mineralcontent of the leaves, while the other 15 samples were used for theenzymic dissolution of the leaves in different fractions,

    From the same sites where the leaf samples were collected, in1980 soil samples were taken underneath 10 chlorotic and 10 healthyvine plants, respectively. Each 10 subsamples were combined to onesample. On the sites 'SieBheim' and 'Iphofen1, soil samples weretaken from the depth 0 to 40 cm and 40 to 80 cm, while in (N1erste1n(samples were taken from the depth 0 to 20 cm and 20 to 60 cm.

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  • IRON CHLOROSIS 717

    Available soil P and K were analyzed by the CAL method, thecommon method used in Germany for the estimation of available P andK. The extractant is consisting of a mixture of Ca lactate, NH4lactate and acetate, pH 4,1 (Schuller^). Iron, Mn, Zn, and Cuwere extracted by DTPA (Lindsay and Norvell9), Carbonate was anal-yzed according to the method of Scheibler^-5 and HCQ3" according tothe method of B ^

    The mineral content of the leaves was analyzed after wet ashingof the ground leaf samples by atomic absorption. Phosphate was de-termined according to the vanadate-molybdate method. For the deter-mination of HC1 soluble Fe 0.5 g of the dried and ground leaf mate-rial was extracted with 20 ml of 0.5 N HC1 and 1.0 N HC1, respec-tively, for 30 min, so that two different HC1 soluble fractions wereobtained, one by the extraction with 0.5 N and one by the extractionwith 1.0 N HC1. Iron was determined by atomic absorption.

    Enzymic Dissolution of_ the Cells

    For the separation of the cells two samples (= 30 leaves total-ly) were combined and thawed slowly in a refrigerator. Then themiddle of the five main leaf veins was isolated. In this fraction,which is called 'leaf veins', Fe, Mn, Zn, Cu, Ca, and P were deter-mined. The rest of the leaves was cut into small pieces and trans-ferred into 10 Erlenmeyer flasks. Then 20 ml of solution No, 1 wasadded.

    Soln. 1: 1% mazerozyme (mixture of pectinase, hemicellulaseand cellulase)

    2% polyethyleneglycoll 400020 mmolar HEPES buffer0.65 molar mannitpH 5.8

    The addition of the buffer was necessary to stabilize the pH,which declined about 0.5 pH units in 2 hours. After 2 h the pH wasreadjusted to pH 5.8 by the addition of 1 N NaOH. The samples wereshaken for 4 hours in a waterbath at 25C and afterwards poured on a100 urn sieve. After washing with 0.65 M mannit the filtrate wascentrifuged. The sediment consisted mainly of intercostal cells.This fraction, called 'intercostal cells' in the following was ana-lyzed for Fe, Mn, Zn, Cu, Ca, and P. The fraction, which did notpass the sieve, was transferred to Erlenmeyer flasks. To this frac-tion 20 ml of solution No. 2 was added.

    Soln. 2: 1% cellulase Onozuka SS*2% polyethyleneglycoll 4Q0Q20% mmolar HEPES buffer0.3 molar mannitpH 5.2

    *Firma Welding and Co., Hamburg, West Germany

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  • 718 MENGEL, BUBL, AND SCHEEER

    The samples were shaken 1n a water bath at 36C for 4 hours andthen poured on a 100 urn sieve. The fraction, which passed throughthe sieve, was centrifuged for 10 minutes at 1000 g. The supertnat-ant in the centrifuge tubes was discharged. After adding 20 ml ofsolution No. 2 to the sediment the centrifuge tubes were shaken for1 h at 36C in a waterbath. By this procedure cells are dissolvedenzymically, whereas the chlorbplasts are not attacked CJacobi6).Chloroplasts were obtained by centrifugation. In the chloroplastfraction Fe, Mn, Zn, Cu, Ca, and P were analyzed. The whole pro-cedure has been described in more detail by bi^

    RESULTS

    Table 1 is showing the Fe concentration in the various chemicaland anatomical fractions of green and chlorotic leaves. Total ironcontent in chlorotic leaves was as high or even higher than 1n greenleaves; however, the amount of HC1 soluble Fe was higher in the

    TABLE 1

    Fe c o n c e n t r a t i o n s in the v a r i o u s and ana tomica l f r a c t i o n s o f green

    and c h l o r o t i c l e a v e s .

    SpieBhein 1979 SpieBheim 1980 Iphofen 1980 Hierstein 1980green ch lorot . green ch lorot . green ch loro t . green ch lo ro t .

    Total Fe 76 89(n = 5)0.5 N HC1 20 18so l . Fe(n = 5)1 N HC1 22 15*so l . Fe(n = 5)Leaf veins(n = 7) ,Intercostal *ce l ls (n = 14) 257 214

    288 290

    ppn Fe, d.m.

    65 64

    30 26*

    37 31*

    58 48

    200 139*538 559Chloroplasts

    (n = 7)Comparison between green and chlorot ic leaves:

    * p = 5 %, ** p = 1' %, * * * p = 0.1 %

    94 97 112 139

    37 33 35 32

    ** *

    40 36 40 37

    72 77 57 60

    203 192 331 233

    1058 1040 726 619

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  • IRON CHLOROSIS 719

    green leaves as compared with the chlorotic ones. The differencesbetween 1 N HC1 soluble Fe of chlorotic and green leaves were sig-nificant for all samples. The amount of Fe extracted by 0.5 N HC1was nearly as high as that extracted by 1 N HC1. The differencesbetween the Fe contents in the veins of healthy and chlorotic leaveswere small and not significant. The Fe content in the intercostalcells of green leaves was significantly higher than in the inter-costal cells of chlorotic leaves. The Fe contents of the chloro-plasts differed very much between the various sites. There were nomajor differences between the Fe content in chloroplasts of greenand chlorotic leaves. However, the chlorotic leaves contained lesschloroplasts.

    As can be seen from Table 2, P concentrations in whole leaveswere significantly higher in the chlorotic samples as compared withthe green ones. The same was true for the P concentrations in theintercostal cells. Also, the Ca concentrations in the chloroticintercostal cells were significantly higher than in the green inter-costal cells. Concentrations of Mn, Zn, and Cu were mainly higherin chlorotic samples than in green samples (Table 3).

    TABLE 2

    Ca- and P concentrat ions in leaves and l e a f - f r a c t i o n s of green andch lo ro t i c leaves.

    SpieBheim 1979 SpieBheim I960 Iphofen 1980 Iphofen 1980green chlorot, green chlorot. green chlorot. green chlorot

    Whole leaf

    Leaf veins

    IntercostalcellsChloroplasts

    Whole leaf

    Leaf veins

    Intercostalcells

    1.57 1.51

    0.69

    1.01

    0.22

    1.7C

    0.93

    0.36*

    0.13 0.24

    % Ca, d.m.

    1.16 1.12

    1.79 1.67

    0.60 1.34

    1.25 1.36% P, d.m.

    0.20 0.30***

    0.11 0.15

    0.09 0.17

    Chloroplasts 0.85 0.71 C.59 0.56

    Comparison between green and.chlorotic leaves:* p = 5 %, ** p = 1 %, *** p = 0.1 %

    1.30 1.50

    1.04 1.23*

    1.04 1.98*

    1.17 1.63

    0.19 0.34

    0.14 0.17*

    0.11 0.16*

    0.13 0.15

    1.32 1.46

    1.43 1.60*

    0.76 1.46*

    0.99 0.84

    0.25 0.38

    0.13 0.20*

    0.15 0.25

    0.35 0.40

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  • 720 MENGEL, BUBL, AND SCHERER

    TABLE 3

    Mn-, Zn- and Cu concentrations in leaves and leaf-fractions of

    green and chlorotic leaves.

    Whole leaf

    Leaf veins

    Intercostalcells

    Chloroplasts

    Whole leaf

    Leaf veins

    Intercostalcells

    Chloroplasts

    Whole leaf

    Leaf veinsIntercostalcells

    SpieBheim 1979green chlorot.

    104

    -

    16

    80

    50

    -

    196

    -

    7

    21

    Chloroplasts 61Cnmparison between* **

    p = 5 %, P = 1

    ***55

    -

    24*

    65

    ***138

    -

    **301

    -

    ***11

    21

    58green arid

    ***

    %, P

    SpieGheim 1980green chlorot.

    ppm Mn

    140

    98

    190

    , d.m.**

    192

    113**

    140

    2444 2870

    ppm Zn

    69

    72

    91

    341

    ppm Cu

    810

    16

    96chlorotic

    = 0.1 %

    , d.m.***

    139

    74

    65*

    297

    , d.m.***

    15

    14***

    24

    99leaves:

    Iphofen 1980green chlorot.

    44

    33

    13

    47

    84

    72

    79

    698

    13

    12

    17

    76

    **57

    30

    8

    31

    ***137

    ***

    112

    88

    796

    12

    11***

    23

    69

    Iphofen 1980green chlorot.

    144

    107

    191

    2359

    55

    60

    -

    -

    5

    9

    16

    86

    ***195

    ***154

    ***127

    1973

    **80**

    68

    -

    -

    7*

    10***

    20

    79

    The most important soil characteristics and nutrient contentsof the soil samples, which were taken underneath green and chloroticvine plants, are shown in Table 4. There was no relationship be-tween the appearance of chlorosis and the pH, the carbonate content,and the contents of CAL-soluble P and K. Also the Fe, Mn, Zn, andCu content and the HCO3" concentration was not related to the chlo-rosis. However, on each of the three sites the clay contents werehigher underneath the chlorotic vine plants.

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  • T A B L E 4N u t r i e n t c o n t e n t s and s o i l c h a r a c t e r i s t i c s o f soi l s a m p l e s t a k e n u n d e r c h l o r o t i c and h e a l t h yv i n e p l a n t s . W i t h t h e e x c e p t i o n of H C O 3 " , t h e d a t a r e l a t e t o a i r d r i e d s o i l . S o i l s a m p l e st a k e n 1 9 7 9 w e r e n o t a n a l y z e d f o r h e a v y rietals.

    site

    SpieR-heim1979

    SpieB-heim1980

    Ip-hofen1980

    Mier-stein1980

    soil depth(cm)

    0-40

    40-80

    0-40

    40-80

    0-40

    40-80

    0-20

    20-60

    symptoms

    green

    chlorot.

    green

    chlorot.

    green

    chlorot.

    green

    chlorot.

    green

    chlorot.

    green

    chlorot.

    green

    chlorot.

    green

    chlorot.

    PH

    7.57.6

    7.67.6

    7.47.4

    7.57.5

    7.37.3

    7.37.3

    7.77.87.77.8

    HC03"

    ppm

    418392

    464493

    507579

    610625

    496524

    502532

    299293293299

    P205mg/

    100 g

    40.827.8

    11.35.7

    32.4

    30.2

    11.7

    7.0

    28.0

    40.0

    19.9

    30.7

    62.3

    91.7

    37.3

    35.0

    K20 mg/

    100 g

    89.661.4

    58.1

    32.0

    73.2

    60.7

    52.4

    35.3

    42.2

    55.7

    26.7

    35.6

    68.9

    62.3

    48.0

    39.0

    Carbonate

    %

    26.3

    26.0

    29.830.0

    25.5

    25.7

    28.5

    28.9

    19.2

    18.8

    22.5

    21.4

    10.3

    10.0

    11.3

    9.8

    Fe

    PPm

    -

    -

    -

    -

    1010

    88

    87

    910

    1010108

    Mn

    ppm

    -

    -

    -

    -

    2324

    2422

    912

    911

    9999

    Zn

    ppm

    -

    -

    -

    -

    77

    94

    914

    88

    301144

    Cu

    ppm

    -

    -

    -

    -

    3329

    1212

    713

    1319

    292477

    clay

    %

    -

    -

    -

    -

    38.746.2

    45.052.4

    29.7

    34.2

    31.1

    34.7

    31.0

    40.933.7

    34.9

    C/3

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  • 722 MENGEL, BUBL, AND SCHERER

    DISCUSSION

    In carbonate soils high HCO3" concentrations may occur CMengelet al.l') due to the high pH level and the dissolution of carbonates.Chlorosis is then likely to appear. This observation was made byBoxma2 for fruit trees. In our investigations all soils were richin carbonate and the pH was higher than 7. But although we foundhigh HCO3" concentrations on all the sites, there was no relation-ship between iron chlorosis and the HCO3" content. However, meas-uring the HCO3" concentration in a soil sample does not provide areliable information about the concentration of HCO3" in the rhizo-sphere and the HC03" uptake of plants. Because of the excretion ofCO2 and H by roots (Mengel and Malissiovasll), the HCO3" concen-tration in the rhizosphere of carbonate soils may differ consider-ably from the HCO3" concentration of the bulked soil. Enhanced C02production in the vicinity of the roots may lead to high HCO3- con-centrations in the rhizosphere.

    Especially HC03" can be accumulated under high soil moistureconditions on soils with a heavy texture. This holds true for ourinvestigations. Higher clay contents were found under chloroticvine plants than under green vine plants. These findings are con-sistent with results of Carter5 who also observed chlorosis on soilswith a high clay content.

    Investigations of Bubl^ with H C03 have shown that tPie rootsof vine plants take up HCO3-. It 1s therefore supposed that on cal-careous soil HCO3" is absorbed by vine plants and that HCQ3" directlyor indirectly affects the Fe transport into the intercostal cells.A similar effect has been observed with nitrate nutrition CMengeland Malissiovas^0). Probably an alkaline nutrition has a detri-mental influence on Fe mobility in vine leaves.

    The reason why HCO3" is hindering the Fe transport into theintercostal cells is not yet clear. Although the total Fe contentof the chlorotic leaves was at least as high as in green leaves oreven higher. HC1 soluble Fe was significantly lower in chloroticleaves as compared to green leaves. This confirms results of Jacob-son' and Mengel et a.1'.

    There was no relationship between the CAL - P content of thesoils and the P content of the leaf samples. But on all sites theP content in the whole leaves and in the intercostal cells, respec-tively, was significantly higher in chlorotic leaves than In thecorresponding samples from green plants. It 1s thus suggested thatFe chlorosis on calcareous soils is not caused by high soil P con-tents. The high P contents 1n the chlorotic leaves are rather theresult of a physiological Fe deficiency. The same holds true forthe high Mn, Zn, and Cu contents of the chlorotic leaves. Also,Bucher* and B00B et al.l found higher P contents in chloroticleaves. Our conclusions that the high P contents 1n chlorotic

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  • IRON CHLOROSIS 723

    leaves are the result and not the cause of the Fe chlorosis areconsistent with the results of Mullner^ and Mengel et al .12,

    REFERENCES

    1. BooB, A., H. Kolesch, and W. Hofner, 1982. Chlorose-Ursachenbe1 Reben (Vitis vinifera L.) am naturlichen Standort. Z,Pflanzenernahr. Bodenkde, 145;246-260.

    2. Boxma, R. 1972. Bicarbonate as the most important soil factorin lime-induced chlorosis in the Netherlands. Plant and Soil37:233-249.

    3. Bubl, W. 1981. Eisen-Chlorose bei der Weinrebe - Loslichkeitund Verteilung von Eisen in grunen und chlorotischen Blatternsowie die Bedeutung des Bicarbonates. Ph. D. Thesis, Fach-bereich 19, Fac. Nutritional Sci., Justus Liebig University,Giessen, West Germany.

    4. Bucher, R. 1976. Der EinfluB hoher Phosphatgaben im Carbonat-boden auf die Aufnahme einiger fur die Rebe wichtigen Spuren-elemente. Weinberg und Keller 23:257-263.

    5. Carter, M. R. 1980. Association of cation and organic anionaccumulation with iron chlorosis of scots pine on parie soils.Plant and Soil 56:293-300.

    6. Jacobi, G. 1974. Biochemische Cytologie der Pflanzenzelle.Ein Praktikum. G. Thieme-Verlag, Stuttgart.

    7. Jacobson, L. 1945. Iron in the leaves and chloroplasts of someplants in relation to their chlorophyll content. Plant Physi-ol. 20:233-245.

    8. Kovanci, I., H. Hakerlerler, and W. Hofner. 1978. Ursachender Chlorosen an Mandarinen (Citrus reticulata bianco) deragaischen Region. Plant and Soil 50:193-205.

    9. Lindsay, W. L. and W. A. Norvell. 1978. Development of a DTPAtest for zinc, iron, manganese and copper. Soil Sci. Soc.Amer. J. 42:421-428.

    10. Mengel, K. and N. Malissiovas. 1981. Bicarbonat als auslosenderFaktor der Eisenschlorose bei der Weinrebe (Vitis vinifera).Vitis 20:235-243.

    11. Mengel, K. and N. Malissiovas. 1982. Light dependent protonexcretion by roots of entire vine plants (Vitis vinifera L.).Z. Pflanzenernahr. Bodenkde. 145:261-267.

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  • 724 MENGEL, BUBL, AND SCHERER

    12. Mengel, K., H. W. Scherer, and N. Malisstovas. 1979. DieChlorose aus der Sicht der Bodenchemie und Rebenernahrung.Matt. Klosterneuburg 29:151-156.

    13. Mullner, L. 1979. Ergebnisse eines Chloroseforschungsprojektes,Mitt. Klosterneuburg 29:141-150.

    14. Rutland, R. B. and M. J. Bukovac, 1971, The effect of calciumbicarbonate on iron absorption and distribution by Chrysanthe-mum morifolium (Ram,). Plant and Soil 35:225-236,

    15. Scheibler, C. 1960. Gasvolumetrische Bestimmung der Kohlenrsaure. In: K. Nehring: Agrikulturchemische Untersuchungs-methoden fur Dunge- und Futternvittel, Boden und Milch, Parey-Verlag, Hamburg und Berlin.

    16. Schuller, H. 1969. Die CAL-Methode, eine neue Methode zurBestimmung des pflanzenverfugbaren Phosphates in Boden. Z.Pflanzenernahr. Bodenkde. 123:48-63.

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