Shifts in xylem vessel diameter and embolisms in grafted apple trees of differing rootstock growth potential in response to drought

We investigated responses of plant growth rate, hydraulic resistance, and xylem cavitation in scion-rootstock-combinations of Malus domestica L. cv. Honeycrisp scions grafted onto a high-shoot vigor (HSV) rootstock, (semi-dwarfing Malling111), or onto a low-shoot vigor (LSV) rootstock, (dwarfing Budagovsky 9), in response to substrate moisture limitation. Adjustments in xylem vessel diameter and frequency were related to hydraulic resistance measurements for high- versus low- vigor apple trees. We observed a greater tolerance to water deficit in the high-shoot compared to the low-shoot vigor plants under water deficit as evidenced by increased growth in several plant organs, and greater scion anatomical response to limited water availability with ca. 25% increased vessel frequency and ca. 28% narrower current season xylem ring width. Whereas water limitation resulted in greater graft union hydraulic resistance of high-shoot vigor trees, the opposite was true when water was not limiting. The graft union of the low-shoot vigor rootstock exhibited higher hydraulic resistance under well-watered conditions. Scions of high-shoot vigor rootstocks had fewer embolisms at low plant water status compared to scions of low-shoot vigor rootstocks, presumably as a result of large differences in xylem vessel diameter. Our results demonstrated that anatomical differences were related to shifts in hydraulic conductivity and cavitation events, a direct result of grafting, under limited soil water.     

Hydraulic resistance components of mature apple trees on rootstocks of different vigours

Dwarfing of fruit trees is often achieved through the use of dwarfing rootstocks. Dwarf trees are characterized by sustained reductions in vegetative growth during the lifetime of the tree. The dwarfing mechanism is not well understood, but it has been hypothesized that hydraulic properties of the rootstock and the graft union are involved. It is hypothesized here that leaf- or stem-specific resistance of at least one hydraulic component of the water transport system would be negatively correlated with rootstock 'vigour', and this could be useful for selection of rootstocks. Hydraulic resistance (R) of fully grown apple trees on a variety of rootstocks of different 'vigours' was measured. Most measurements were with the evaporative flux (EF) method, where water uptake measured with sap flow sensors was related to the pressure gradient from soil (taken as pre-dawn leaf) and midday root (taken as covered root-sucker), stem (from covered leaf), and exposed and shaded leaf water potentials (Psi(l)). R of trees on dwarfing M9 rootstock was compared with that of more vigorous MM106 and MM111 rootstocks in Israel and Vermont, USA. In Israel, M9 consistently had higher leaf-specific hydraulic resistance (R(l)) in the soil to scion stem pathway, but this difference was only significant for one summer. R was larger in M9 between the root and stem, implicating the graft union as the site of increased resistance. In Vermont, R(l) of 9- and 10-year-old trees on six rootstocks of various vigours was not consistently related to vigour, and stem-specific resistance (R(s)) increased with increasing vigour. High pressure flow meter (HPFM) measurements gave a lower R than the EF method in all but one case, perhaps indicating a significant amount of xylem dysfunction in these trees, and demonstrated the increased resistivity of stem sections that included dwarf graft unions as compared with non-graft stem sections. It is concluded that stem- and leaf-specific R are not consistently positively correlated with dwarfing, although the increased resistivity of the graft union in dwarfing rootstocks may influence the transport of water and other elements across the graft union, and therefore be involved in the dwarfing mechanism.     

Rootstock‐to‐scion transfer of transgene‐derived small interfering RNAs and their effect on virus resistance in nontransgenic sweet cherry

Small interfering RNAs (siRNAs) are silencing signals in plants. Virus‐resistant transgenic rootstocks developed through siRNA‐mediated gene silencing may enhance virus resistance of nontransgenic scions via siRNAs transported from the transgenic rootstocks. However, convincing evidence of rootstock‐to‐scion movement of siRNAs of exogenous genes in woody plants is still lacking. To determine whether exogenous siRNAs can be transferred, nontransgenic sweet cherry (scions) was grafted on transgenic cherry rootstocks (TRs), which was transformed with an RNA interference (RNAi) vector expressing short hairpin RNAs of the genomic RNA3 of Prunus necrotic ringspot virus (PNRSV‐hpRNA). Small RNA sequencing was conducted using bud tissues of TRs and those of grafted (rootstock/scion) trees, locating at about 1.2 m above the graft unions. Comparison of the siRNA profiles revealed that the PNRSV‐hpRNA was efficient in producing siRNAs and eliminating PNRSV in the TRs. Furthermore, our study confirmed, for the first time, the long‐distance (1.2 m) transfer of PNRSV‐hpRNA‐derived siRNAs from the transgenic rootstock to the nontransgenic scion in woody plants. Inoculation of nontransgenic scions with PNRSV revealed that the transferred siRNAs enhanced PNRSV resistance of the scions grafted on the TRs. Collectively, these findings provide the foundation for ‘using transgenic rootstocks to produce products of nontransgenic scions in fruit trees'.     

Etiology of three recent diseases of citrus in São Paulo State: sudden death, variegated chlorosis and huanglongbing

The state of S?o Paulo (SSP) is the first sweet orange growing region in the world. Yet, the SSP citrus industry has been, and still is, under constant attack from various diseases. In the 1940s, tristeza-quick decline (T-QD) was responsible for the death of 9 million trees in SSP. The causal agent was a new virus, citrus tristeza virus (CTV). The virus was efficiently spread by aphid vectors, and killed most of the trees grafted on sour orange rootstock. Control of the disease resided in replacing sour orange by alternative rootstocks giving tolerant combinations with scions such as sweet orange. Because of its drought resistance, Rangpur lime became the favourite alternative rootstock, and, by 1995, 85% of the SSP sweet orange trees were grafted on this rootstock. Therefore, when in 1999, many trees grafted on Rangpur lime started to decline and suddenly died, the spectre of T-QD seemed to hang over SSP again. By 2003, the total number of dead or affected trees was estimated to be over one million. The new disease, citrus sudden death (CSD), resembles T-QD in several aspects. The two diseases have almost the same symptoms, they spread in time and space in a manner strikingly similar, and the pathological anatomy of the bark at the bud union is alike. Transmission of the CSD agent by graft-inoculation has been obtained with budwood inoculum taken not only on CSD-affected trees (grafted on Rangpur lime), but also on symptomless trees (grafted on Cleopatra mandarin) from the same citrus block. This result shows that symptomless trees on Cleopatra mandarin are tolerant to the CSD agent. Trees on rootstocks such as Sunki mandarin or Swingle citrumelo are also tolerant. Thus, in the CSD-affected region, control consists in replacing Rangpur lime with compatible rootstocks, or in approach-grafting compatible rootstock seedlings to the scions of trees on Rangpur lime (inarching). More than 5 million trees have been inarched in this way. A new disease of sweet orange, citrus variegated chlorosis (CVC), was observed in 1987 in the Triangulo Mineiro of Minas Gerais State and the northern and north-eastern parts of SSP. By 2000, the disease affected already 34% of the 200 million sweet orange trees in SSP. By 2005, the percentage had increased to 43%, and CVC was present in all citrus growing regions of Brazil. Electron microscopy showed that xylem-limited bacteria were present in all symptomatic sweet orange leaves and fruit tissues tested, but not in similar materials from healthy, symptomless trees. Bacteria were consistently cultured from twigs of CVC-affected sweet orange trees but not from twigs of healthy trees. Serological analyses showed the CVC bacterium to be a strain of Xylella fastidiosa. The disease could be reproduced and Koch's postulates fulfilled, by mechanically inoculating a pure culture of X. fastidiosa isolate 8.1.b into sweet orange seedlings. The genome of a CVC strain of X. fastidiosa was sequenced in SSP in the frame of a project supported by FAPESP and Fundecitrus. X. fastidiosa is the first plant pathogenic bacterium, the genome of which has been sequenced. Until recently, America was free of huanglongbing (HLB), but in March 2004 and August 2005, symptoms of the disease were recognized, respectively in the State of S?o Paulo (SSP) and in Florida, USA. HLB was known in China since 1870 and in South Africa since 1928. Because of its destructiveness and its rapid spread by efficient psyllid insect-vectors, HLB is probably the most serious citrus disease. HLB is caused by a phloem sieve tube-restricted Gram negative bacterium, not yet available in culture. In the 1990s, the bacterium was characterized by molecular techniques as a member of the alpha proteobacteria designated Candidatus Liberibacter africanus for the disease in Africa, and Candidatus Liberibacter asiaticus for HLB in Asia. In SSP, Ca. L. asiaticus is also present, but most of the trees are infected with a new species, Candidatus Liberibacter americanus.     

Variation in the content of bioflavonoids of orange as affected by scion,rootstock, and fruit part

Fruits of citrus cultivars contain bioflavonoids and some other important secondary metabolites in pharmaceutical and nutritional industries. The present experiment was designed to investigate the correlation between the content of flavonoid components like naringin, hesperidin, and neohesperidin of the scions fruits and the same parameters in rootstocks fruits. Six-year-old trees including four citrus cultivars of ‘Moro’, ‘Mars’, ‘Salustiana’, and ‘Italian’ which were previously grafted on the four different rootstocks including ‘Yuzu’, ‘Shelmahalleh’, ‘Citromelo’, and ‘Sour orange’ were selected as experimental trees. The content of the mentioned flavonoids was investigated in the peel and pulp of the fruits of both scions and rootstocks. The results showed that the measured parameters were significantly influenced by scions, rootstocks, and tissues. Based on the obtained results, it can be suggested that the accumulation of chemicals in citrus fruit depends on genetic and inherent abilities of the scion, more than what was previously believed, while the rootstock can also play an important role in the accumulation of these substances.     

Phytophthora collar rot of apple: influence of the rootstock on scion variety resistance

In field experiments with young trees great differences were found in the resistance to Phytophthora cactorum of Cox's Orange Pippin apple scions grafted on different clonal rootstocks. The rootstock effect on scion resistance was inversely related to the effect on tree vigour: the rootstocks inducing high resistance were dwarfing (M. 9) or semi-dwarfing (M. 7, M. 26, MM. 106), and those inducing low resistance were vigorous or very vigorous (M. 25, MM. 104, MM. 109). Mean lesion lengths in Cox on MM. 104 were five to eight times greater than those in Cox on M. 9. The rootstock influence on scion resistance was associated primarily with effects on the rate of lesion extension: during the early stages only of host colonization there appeared to be threshold extension rates below which host resistance factors effectively suppressed a large proportion of infections. The influence of the root-stock on scion resistance was apparently unrelated to inherent rootstock resistance. On all rootstocks Cox showed diminished resistance to infection during the period from the swelling of buds to the early stages of shoot growth. Although most susceptible during the ‘mouse-ear’ and ‘pink bud’ stages of development, suscpetibility was not associated with flowering per se. Rootstock type did not affect the resistance of Cox scions to P. syringae, for which the period of susceptibility to infection occurred in the dormant season.     

Effect of rootstock on apple (Malus domestica) tree water relations

The effects of rootstock on mid-season water relations, under orchard conditions of non-limiting soil moisture, were determined for bearing 'Empire' apple trees ( Malus domestica Borkh.) on the clonal rootstocks M9, M26, M7, MM106, and MM104 (most to least dwarfing) in their sixth and seventh growing seasons. Stem water potentials (ψ^stem) of trees on M9 and M26 were more negative at midday, under warm, sunny conditions, than were the trees on the other three rootstocks. However, change in ψ^stem per change in stem distance through the canopy (water potential gradient) did not vary among rootstocks at midday. There was no rootstock effect on diurnal variation in transpiration or stomatal conductance. Differences in water storage capacitance, relative to tree size, were determined in a separate study but did not account for the differences observed in ψ^stem. Calculated hydraulic conductivities of xylem water transport suggest that rootstocks differ in their ability to conduct water to the scion, but hydraulic conductivity of the scion was not affected by rootstock. Root-stock differences in hydraulic conductivity were not accounted for by differences in tree size.     

Rootstock effects on xylem conduit dimensions and vulnerability to cavitation of <Emphasis Type="Italic">Olea europaea</Emphasis> L.

Two clones of Olea europaea L. were studied for their potential impact on hydraulic architecture and vulnerability to xylem cavitation, when used as rootstocks. The clones used were “Leccino Minerva” (LM), showing vigorous growth and “Leccino Dwarf” (LD) with strongly reduced growth. Self-rooted LM and LD plants as well as their grafting combinations were compared, namely, LM/LD (Leccino Minerva grafted onto Leccino Dwarf rootstock) and LD/LM (Leccino Dwarf grafted onto Leccino Minerva rootstocks). Plants with LD roots (LD and LM/LD) showed significantly reduced leaf surface area compared with plants with LM roots. Xylem conduits of LD shoots were 25% more numerous than in LM shoots. When grafted onto LM rootstocks, however, LD shoots produced consistently wider and longer vessels than measured in LD self-rooted plants. This caused LD/LM plants to increase stem vulnerability to cavitation with threshold pressures for cavitation (P _c) of less than 0.5 MPa compared with LD self-rooted plants that had P _c of over 2.0 MPa. By contrast, although LD rootstocks caused some reduction of vessel diameter and length of LM scions, their influence on LM hydraulic architecture was too small to reduce vulnerability to cavitation of LM scions with respect to that measured for LM self-rooted plants. Our conclusion is that although dwarfing rootstocks effectively reduce grafted plant size, they do not necessarily confer higher resistance to xylem cavitation to scions which would improve plant resistance to drought.     

The effect of root pressurization on water relations, shoot growth, and leaf gas exchange of peach (Prunus persica) trees on rootstocks with differing growth potential and hydraulic conductance

It is well known that rootstocks can have an effect on the vegetative growth and development of the tree; however, there has been no clear explanation about the physiological mechanism involved in this phenomenon. Evidence indicates that the rootstock effects on tree vegetative growth may be related to hydraulic limitations of the rootstock. The objective of these experiments was to investigate the shoot growth, water potential, and gas exchange of peach trees on different rootstocks in response to manipulations of water relations of trees on rootstocks that differ in root hydraulic conductance. Tree water relations were manipulated by applying different amounts of pneumatic pressure on the root system and then relative shoot extension growth rate, tree transpiration rate, leaf water potential, leaf conductance, leaf transpiration, and net CO(2) exchange rate responses were measured. Root pressurization increased leaf water potential, relative shoot extension growth rate, leaf conductance, leaf transpiration, and net CO(2) exchange rates of trees on both vigorous and dwarfing rootstocks. There was a significant positive linear correlation between applied pneumatic pressure and tree transpiration rate and leaf water potential. Leaf conductance, transpiration rate, and net CO(2) exchange rate as well as relative shoot extension growth rates were also positively correlated with the applied pneumatic pressure on the root system. These relationships were consistent across both vigorous and size-controlling rootstocks, indicating that rootstock hydraulic limitation may be directly involved in the vegetative growth control of peach trees.     

Root and stem hydraulic conductivity as determinants of growth potential in grafted trees of apple (Malus pumila Mill.)

The anatomy of the graft tissue between a rootstock and its shoot (scion) can provide a mechanistic explanation of the way dwarfing Malus rootstocks reduce shoot growth. Considerable xylem tissue disorganization may result in graft tissue having a low hydraulic conductivity (k(h)), relative to the scion stem. The graft may influence the movement of substances in the xylem such as ions, water and plant-growth-regulating hormones. Measurements were made on 3-year-old apple trees with a low-pressure flow system to determine k(h) of root and scion stem sections incorporating the graft tissue. A range of rootstocks was examined, with different abilities of dwarfing; both ungrafted and grafted with the same scion shoot cultivar. The results showed that the hydraulic conductivity (k(hroot)) of roots from dwarfing rootstocks was lower compared with semi-vigorous rootstocks, at least for the size class of root measured (1.5 mm diameter). Scion hydraulic conductivity (k(hs)) was linked to leaf area and also to the rootstock on to which it was grafted, i.e. hydraulic conductivity was greater for the scion stem on the semi-vigorous rootstock. Expressing conductivities relative to xylem cross-sectional areas (k(s)) did not remove these differences suggesting that there were anatomical changes induced by the rootstock. The calculated hydraulic conductivity of the graft tissue was found to be lower for grafted trees on dwarfing rootstocks compared to invigorating rootstocks. These observations are discussed in relation to the mechanism(s) by which rootstock influences shoot growth in grafted trees.     

Relationships between xylem vessel characteristics, calculated axial hydraulic conductance and size-controlling capacity of peach rootstocks

Background and Aims

Previous studies indicate that the size-controlling capacity of peach rootstocks is associated with reductions of scion water potential during mid-day that are caused by the reduced hydraulic conductance of the rootstock. Thus, shoot growth appears to be reduced by decreases in stem water potential. The aim of this study was to investigate the mechanism of reduced hydraulic conductance in size-controlling peach rootstocks.

Methods

Anatomical measurements (diameter and frequency) of xylem vessels were determined in shoots, trunks and roots of three contrasting peach rootstocks grown as trees, each with different size-controlling characteristics: ‘Nemaguard’ (vigorous), ‘P30-135’ (intermediate vigour) and ‘K146-43’ (substantially dwarfing). Based on anatomical measurements, the theoretical axial xylem conductance of each tissue type and rootstock genotype was calculated via the Poiseuille–Hagen law.

Key Results

Larger vessel dimensions were found in the vigorous rootstock (‘Nemaguard’) than in the most dwarfing one (‘K146-43’) whereas vessels of ‘P30-135’ had intermediate dimensions. The density of vessels per xylem area in ‘Nemaguard’ was also less than in ‘P30-135’and ‘K146-43’. These characteristics resulted in different estimated hydraulic conductance among rootstocks: ‘Nemaguard’ had higher theoretical values followed by ‘P30-135’ and ‘K146-43’.

Conclusions

These data indicate that phenotypic differences in xylem anatomical characteristics of rootstock genotypes appear to influence hydraulic conductance capacity directly, and therefore may be the main determinant of dwarfing in these peach rootstocks.Key words: Prunus, rootstock, vessel diameter, hydraulic conductance, dwarfing, xylem anatomy, Poiseuille–Hagen     

The Anatomy of the Developing Bud Union and its Relationship to Dwarfing in Apple

Light microscopy has been used to study the effect of dwarfingand semi-dwarfing apple rootstocks on the early developmentof bud-unions with 'Gala', and the anatomy of 2-year-old bud-unionsbetween 'Bramley' and the same rootstocks. The bridging of thecut edges of the cambia of bud and rootstock was achieved bydifferentiation of callus formed at an early stage in budding.New cambial cells were aligned at right angles to the pre-existingcambia, with their long axes horizontal. Subsequently-formedxylem adopted this arrangement, so that fibres and vessels werearranged obliquely to the axis of the stem. At the interfacebetween the bud and dwarfing rootstocks vessels with smallerthan normal diameter were formed, indicating the presence ofelevated levels of auxin in this region. In addition, littlexylem was produced in the adjacent rootstock tissue. In thecase of semi-dwarfing rootstocks, the rootstock produced normalxylem after a brief interruption. We suggest that failure ofauxin to cross the bud-union interface in the case of the dwarfingrootstocks leads to reduced rootstock xylem formation, and hencea poor supply of water and minerals to the scion, and this underliesthe dwarfing effect.Copyright 1994, 1999 Academic Press Apple, budding, dwarfing, anatomy, graft union     

Development,histological observations and Grapevine leafroll‐associated virus‐3 localisation in in vitro grapevine micrografts

Development, histological process and Grapevine leafroll‐associated virus‐3 localisation were studied in micrografts of three scion/rootstock combinations: healthy/healthy, healthy/infected and infected/healthy. Earlier bud break and faster growth in scions of micrografts were obtained when the healthy shoot segments were used as scions, while earlier bud break in rootstocks and greater fresh weight of roots in micrografts were produced when the healthy shoot segments were used as rootstocks. All histological processes including callus initiation and formation in micrografting conjunctions, and initiation of new cambial cells followed by vascular bundle development connecting scions and rootstocks were similar in micrografts, regardless of the sanitary status of the scions and rootstocks used for micrografting. Virus infection in micrografting conjunctions and systematic infection in micrografts were much more efficient and faster in micrografting combination of the infected scions/healthy rootstocks than in the healthy scions/infected rootstocks. To the best of our knowledge, this is the first report addressing histological process of micrograft development and virus localisation in micrografts. In vitro culture system established in this study facilitates studies on the ‘pure’ impact of the viral infection on micrografting.     

Changes in the anatomical structure of tree rings of the rootstock and scion in the heterografts of Siberian pine

The analysis of the anatomical characteristics of tree rings of the Pinus sibirica scions and Pinus sylvestris rootstocks is presented. The main anatomical features maintain the seasonal dynamics characteristic of the reference (ungrafted) trees of scion and rootstock, pointing to the stability of the genetic control of the xylem differentiation. However, various anatomical characteristics are reduced for rootstocks and increased for scions relative to the reference trees. A mechanism consistent with our results is that the graft union zone creates a barrier for the ascending and descending transport of substances, including phytohormones. The reaction of the scion and rootstock to climatic factors is weakened relative to the reaction of the reference trees. Thus, the presumed shift towards phytohormone disbalance reduces the influence of external factors on xylem differentiation. The interannual variation of the individual characteristics of the scions and rootstocks is increased relative to the reference trees, reflecting either the influence of the graft partners on each other or the non-uniform distribution of the growth regulators across the graft union. Thus, heterografts can be used to evaluate the interaction of the internal and external mechanisms of xylem differentiation.     

The relationship of hydraulic conductance to root system characteristics of peach (Prunus persica) rootstocks

Specific rootstocks can differentially influence the vegetative growth and development of fruit trees. However, the physiological mechanism involved in this phenomenon has been elusive. Recent research comparing different peach ( Prunus persica L. Batsch) rootstocks suggests that the rootstock effect on vegetative growth in peach trees is associated to water relations and more specifically to differences in rootstock hydraulic conductance. This study was intended to confirm differences in hydraulic characteristics of similar size peach trees grafted on different rootstocks and to examine root system characteristics that could be associated with rootstock hydraulic limitations. Trees on rootstocks that were known to have a size-controlling effect when grown under field conditions had lower rootstock conductance than trees on the vigorous (control) rootstock when rootstock hydraulic conductance was measured with both the high-pressure and evaporative flow methods. Rootstocks with the lowest hydraulic conductance had less fine root surface area and length per unit root dry weight than the more vigorous (control) rootstock. However, contrary to previous field studies, in this study there were no significant differences in dry matter production and distribution among trees on the different rootstocks suggesting that whatever the normal growth control mechanism was, it did not differentially influence growth under the specific conditions of this study. This research confirmed that peach rootstocks exhibiting size-controlling behavior under field conditions differed in their hydraulic and morphological characteristics under controlled growth conditions even when those growth conditions negated the expression of the size-controlling behavior.     

The resistance of sour orange to Citrus tristeza virus is mediated by both the salicylic acid and RNA silencing defence pathways

Citrus tristeza virus (CTV) induces in the field the decline and death of citrus varieties grafted on sour orange (SO) rootstock, which has forced the use of alternative decline‐tolerant rootstocks in affected countries, despite the highly desirable agronomic features of the SO rootstock. Declining citrus plants display phloem necrosis below the bud union. In addition, SO is minimally susceptible to CTV compared with other citrus varieties, suggesting partial resistance of SO to CTV. Here, by silencing different citrus genes with a Citrus leaf blotch virus‐based vector, we have examined the implication of the RNA silencing and salicylic acid (SA) defence pathways in the resistance of SO to CTV. Silencing of the genes RDR1, NPR1 and DCL2/DCL4, associated with these defence pathways, enhanced virus spread and accumulation in SO plants in comparison with non‐silenced controls, whereas silencing of the genes NPR3/NPR4, associated with the hypersensitive response, produced a slight decrease in CTV accumulation and reduced stunting of SO grafted on CTV‐infected rough lemon plants. We also found that the CTV RNA silencing suppressors p20 and p23 also suppress the SA signalling defence, with the suppressor activity being higher in the most virulent isolates.     

The physiological and nutritional responses of seven different citrus rootstock seedlings to boron deficiency

Key message

Carrizo citrange was the most tolerant citrus rootstock to B-deficiency and some physiological performance could be attributed to the decreased mineral nutrient concentrations caused by B-deficiency.

Abstract

Boron (B) is an essential microelement for normal growth and development in vascular plants, and adequate B nutrition is crucial for agricultural production. Although citrus plants are not classified as the most sensitive species to B-deficiency, the occurrence of B-deficiency has been reported in the major citrus producing countries of the world, including the east and south of China. In this study, in order to evaluate the effects of B-deficiency on plant growth, root-morphological traits, B and other nutritional responses of citrus rootstock and to investigate the relationship between this physiological performance and mineral nutrients seven common rootstock seedlings, including Trifoliate orange (TO), Carrizo citrange (CC), Chongyi tangerine (CT), Red tangerine (RT), Cleopatra mandarin (CM), Fragrant citrus (FC), and Sour orange (SO), were treated by B-deficiency (0 mg L^?1) or moderate B (0.25 mg L^?1). All the seedlings were grown in hydroponics situation with modified 1/2-strength Hoagland’s solution under greenhouse conditions for 10 weeks. The results showed that B-deficiency inhibited the growth and development of all tested citrus rootstocks, but substantial differences were observed among these rootstocks. Different visible symptoms were observed both in the leaf and root. Corking of the leaf veins and leaf yellowing symptoms were observed on all rootstock genotypes except on CC, which exhibited a little discoloration at the end of the experiment. In addition, root growth of the citrus seedlings were also decreased by B-deficiency, but the decreases were more obvious in TO and FC. It was worth noting that B-deficiency inhibited lateral root growth and development more significantly than tap root, but not in lateral root initiation. The different performance of these rootstock genotypes indicated that CC was the most tolerant while TO was the most sensitive to B-deficiency. In addition, under B-deficiency conditions, not only the B concentration, but also the other mineral nutrient concentrations were influenced, especially in Mg, Fe and Mn. This change in nutrient concentrations might partly contribute to the seedlings’ physiological performances under B-deficiency.     

Comparative sensitivity of 'Prior Lisbon' lemon and 'Valencia' orange trees to foliar sodium and chloride concentrations

Abstract While citrus rootstocks differ in capacity for sodium and chloride ion exclusion, citrus scion species also vary in foliar sensitivity to NaCl salinisation. Of two common scions, ‘Lisbon’ lemon appears more sensitive, whereas ‘Valencia’ orange in less sensitive to leaf salt. In an attempt to explain this difference. ‘Valencia’ orange (Citrus sinensis [L.] Osbeck) and ‘Prior Lisbon’ lemon (Citrus limon [L.] Burm. F.) were budded to rootstocks known to differ in their ability to exclude sodium ions viz, the strong excluder Trifoliata (Poncirus trifoliata [L.] Raf.), and the weaker excluder Troyer citrange (C. sinensis×P. trifoliata); neither rootstock shows strong exclusion of chloride ions. Budded trees were held under a photosynthetic photon flux density of 450 μmol m ² S ¹ and watered with nutrient solution containing either 0 or 50 mol m ³ NaCl. Growth and photosynthetic responses were measured over 58 d following onset of salinization: salinity effects on leaf gas exchange were studied in relation to changes in leaf water status, compatible solutes and foliar content of sodium and chloride ions, over that same period. Once root-zone salinization began to influence leaf solutes (day 30 onwards), lemon showed a steeper increase in leaf chloride than occurred for orange. Although rootstock differences were without effect on this ingress of chloride ions for either scion, sodium ions were excluded from both scions to a larger extent by Trifoliata than by Troyer citrange. Carbon dioxide assimilation of scion foliage was reduced earlier and to a much larger extent by rootzone salinization in lemon than in orange. Furthermore, comparisons of CO₂ assimilation in relation to leaf tissue solutes between scions (on either rootstock) showed stronger responses for both sodium and chloride ions in lemon than in orange. Faster ingress of chloride into lemon leaves was identified as the crucial factor which predisposed towards expression of that contrast between scions. Although contrasts between scions in photosynthetic responses to salinization matched a faster ingress of chloride into lemon than into orange leaves, the sharper photosynthetic response of ‘Prior Lisbon’ lemon to salinity was not solely attributable to higher concentrations of chloride ions (cell sap basis). A difference between species in subcellular compartmentation of the chloride ion under saline conditions was invoked.