C A L I F O R N I A

Boron Mobility and Consequent Management in Different Crops

oron deciency and toxicity occur throughout agricultural regions worldwide. To identify and correct an imbalance of B requires knowledge of the processes governing uptake, remobilization and distribution in the plant. Mobile or Not Mobile in Plant Tissue? Boron is now known to be mobile in the phloem of all species that utilize polyols (complex sugars) as a primary photosynthetic metabolite. In these species a polyol-B-polyol complex is formed in the photosynthetic tissues and is transported in the phloem to currently active sink regions such as vegetative or reproductive meristems. In species that do not produce signicant quantities of polyols, B once delivered to the leaf in the transpiration stream cannot reenter the phloem, resulting TABLE 1. in essentially complete phloem Organ immobility. In species for Leaf which B is immoHull Shell bile, B moves with Kernel the transpiration
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stream and once it enters a leaf it tends to remain. Thus, B will accumulate at the sites of termination of leaf veins. A steep gradient in B concentration has often been found such that B concentration in petioles and midribs < middle of lamina < margins and tips. This principle is illustrated in It has been accepted that Figure 1 in which the disthe uptake of boron (B) is a tribution of B within a passive (non-metabolic) mature leaf of walnut and process and that it is a apple is contrasted. In walphloem immobile element, nut, in which B is immoso once incorporated into bile, the highest B accutissue, it cannot be remobimulation occurred at the lized to supply the needs of tip and margin of the leaf. other plant tissues. Recent
work by Brown and coworkers, however, has now demonstrated that the physiology (mobility) of B varies dramatically between plant species and that our current knowledge of the symptoms and management of B nutrition must be re-examined on a species by species basis.

Boron Distribution in Plant Tissue Figure 1 also illustrates B distribution in apple. In this species leaf B concentrations were signicantly lower than in walnut, and there was very little difference in B accumulation across the leaf.

Boron concentration (ppm dry wt.) in leaf and fruit organs of four tree species.
B mobile Almond 42 170 34 43 Apple 41 51 (peel) 34 (pulp) 54 (core) B immobile Pistachio Walnut 130 33 2 1 295 40 9 4

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This uniform low distribution of B in apple does not correlate with leaf venation pattern and is not consistent with the hypothesis that B distribution is determined solely by transpiration. This same leaf distribution was observed in almond, peach and plum, suggesting that B distribution in these species is not governed by transpiration and is suggestive of phloem B mobility. Evidence of phloem B mobility or immobility can also be discerned from the distribution of B within different organs of a given species. For example, under eld conditions pistachio and walnut both contain the highest B concentration in mature leaves and the lowest B concentration in fruit and seed tissue (Table 1). In contrast, almond or apple grown at the same site had highest B concentration in hull (fruit tissue), with much lower B in the leaves (Table 1). The concentration of B in leaves of different ages within a species can also provide evidence of B mobility. The occurrence of higher B concentrations in old or mature leaves in comparison to younger leaves is evidence of B immobility, while higher B concentrations in younger leaves is an indication of B mobility since these leaves have transpired less water than older leaves. The results in Table 2 suggest that B is phloem immobile in pecan, tomato, strawberry and walnut, while B is phloem mobile in apple, apricot, pear, grape, loquat, peach, celery, olive, and pomegranate. These differences in the site of accumulation of B in tissues will, in turn, determine where within a plant the symptoms of B toxicity will occur. Boron Toxicity Symptoms The difference in B mobility also results in difference in the expression of B toxicity symptoms. Because plants in which B is immobile always accumulate B in the tip and edge of old leaves like walBetter Crops/Vol. 82 (1998, No. 2)

1263 58

44 211 41 725

35

181

30

33

Apple Boron mobile

Walnut (terminal leaflet) Boron immobile

Figure 1. Leaf B concentration (ppm) in eld grown apple and walnut. Leaves were collected at the end of the growing season. The two species were grown in close proximity and received the same irrigation.

nut in Figure 1, B toxicity symptoms in those species are always exhibited as leaf tip and edge burn (Figure 2a, pistachio). On the other hand, for those plants in which B is mobile, instead of the marginal leaf burn, these species exhibit B toxicity as die back in young shoots (Figure 2b, almond), profuse gumming in the leaf axil and the appearance of brown, corky lesions along stems and petioles (Figure 2c, almond). Die back induced by B toxicity has been reported in almond, apple, apricot, cherry, peach, pear, plum, and prune etc. The occurrence of these unusual symptoms of B toxicity are not,

Figure 2a. Boron is immobile in pistachio, so toxicity symptoms appear as leaf tip and edge burn.

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however, restricted solely to the members of the Prunus, Malus and Pyrus genera as discussed earlier. For example, celery responds to B toxicity by producing deformed young leaves, bitter and misshapen stems, while the tip burn symptom is absent. Boron is known to be phloem mobile in celery, since celery like the members of the Prunus genera, utilizes a polyol as a primary transported photosynthate. The two kinds of B toxicity symptoms described above are a consequence of the difference in B mobility. In short, for those species in which B is immobile, B toxicity is exhibited as tip/edge burn of old leaves, while for those in which B is mobile, meristematic dieback is the primary symptom of B toxicity.

Diagnosis and Fertilization In general, for the majority of plant species, B is phloem immobile, however B is phloem mobile in many important crop species such as those presented in Table 2. Preliminary evidence also suggests that many other crop species may exhibit phloem B mobility (i.e. coffee) though this is yet to be veried. The difference in B mobility indicated above also profoundly inuences the diagnosis of plant B status and the correction of deciencies TABLE 2. Leaf B concentration (ppm dry wt.) along a shoot in various and toxicities. plant species. Currently pracSpecies Basal Middle Apical Remarks ticed sampling Pecan 303 119 30 B-immobile techniques and Tomato 721 318 94 B-immobile symptom descripStrawberry 512 176 68 B-immobile tions have been Walnut 304 127 48 B-immobile based on the Apple 50 56 70 B-mobile premise that B is Apricot 45 60 81 B-mobile not mobile in the Pear 42 57 62 B-mobile plant. Selection of Celery 32 49 104 B-mobile tissue samples and Grape 74 55 88 B-mobile determination of Loquat 72 101 162 B-mobile Olive 42 51 56 B-mobile critical nutrient Peach 53 57 208 B-mobile concentrations are Pomegranate 21 20 111 B-mobile all fundamentally
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dependent on the phloem mobility of B. Table 2 illustrates that B does not accumulate in the older leaves of species in which B is mobile. Thus, old leaves are not suitable for determination of B toxicity. Rather young apical leaves or fruit tissue may be a superior indicator in these species (Table 1, 2). This observation has led to the widespread use of hull B as a determinant of B status in almond in California. In species with limited B mobility however, the B concentration in the old leaves (Table 2) remains a good indicator of B toxicity. For the diagnosis of B deciency, the use of a recently matured, or fully expanded leaf is inappropriate if B is phloem immobile since the B concentration of a developed leaf may not reect the B status of growing tissues for which a constant B supply is most critical. This can only be achieved by sampling growing tissues. This is an inherently difcult and inconsistent process. However, it is the only valid approach. By contrast, in species that exhibit mobility, mature leaves are appropriate for assessing B deciency since their content reects the B status of the entire plant, including young growing tissues. In these species B depletion in the soil will not impact meristematic

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growth until the soluble B pool of mature tissues has also been depleted. The management of B fertilization in plants is directly inuenced by patterns of B mobility. Experimental evidence clearly demonstrates that foliar applied B can be retranslocated to the growing organs in species with signicant phloem B mobility. This suggests that foliar B application can be used effectively at any time functional leaves are present to correct B deciency, Figure 2b. (At left) Boron is mobile in almond, and toxicity may and to supply B to future ower cause dieback in young shoots. and fruit tissue. The signicant Figure 2c. (At right) Profuse gumming in the leaf axil and benets of foliar B application appearance of brown, corky lesions along stems on fruit set that has been and petioles are symptoms of B toxicity in almond. observed in many tree species such as almond, plum, prune and others is terize the patterns of B mobility in diverse a consequence of this mobility. In species plant species. However, the following prein which B is immobile, however, foliar dictions can now be made about several applied B cannot be retranslocated from important eld crops. the site of application and cannot supply In corn, wheat, alfalfa and vegthe B requirements of tissue not yet etable crops except those mentioned earformed. Hence, in these species B appli- lier, B is immobile and must, therefore, be cations must be made directly to the tis- supplied at all stages of plant growth. sue of interest. In fruit crops, where B is Foliar application can be used to essential for the owering process, this correct deciency in current tissues but means that B applications can only be will have minimal effect on new plant effectively used if applied directly to the growth. ower buds or owers themselves. In some species, there may be cultivars in which a small amount of mobiliSummary ty may occur. Knowledge of the relative mobility of This may explain the differences in B within a particular species determines sensitivity to B deciency that are occathe approach that should be used to sam- sionally observed between cultivars. ple and diagnose B status. This knowl- Further work on this topic is required. edge also determines the optimum fertilization strategy that should be used and Dr. Brown is Associate Professor and Dr. Hening Hu helps in our understanding of the causes is a Postdoctoral Research Scientist, Department of and consequences of B deciency. Pomology, University of California, Davis. Further work is underway to fully charac-

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