H+/OH- Excretion and Nutrient Uptake in Upper and Lower Parts of Lupin (Lupinus angustifolius L.) Root Systems

The cultivation of narrow-leafed lupins (Lupinus angustifoliusL.) increase rates of subsoil acidification, and this is thoughtto be partly related to their pattern of nutrient uptake andH⁺/OH^- excretion. The main hypothesis of this study was thatH⁺ and OH^- excretion is not distributed evenly over the entirelength of the root system but is limited to zones where excesscation or anion uptake occur. Seedlings of nodulated lupinswere grown in solution culture using vertically split pots thatallowed the upper and lower zones of the root system to be suppliedwith varying concentrations of K⁺ and NO^-₃. Net H⁺/OH^- excretionwas equated to the addition of NaOH/HCl required to maintaina constant pH in the nutrient solution during a 4-d treatmentperiod and nutrient uptake was measured by depletion from solutionin each zone of the split pots. The excess of cation over anion uptake was positively correlatedwith H⁺ excretion in each rooting zone. In zones where K⁺ wassupplied at 1200 µM, cation uptake was dominated by K⁺and up to twice as much H⁺ was excreted than in zones whereK⁺ was absent. In zones where NO^-₃ was supplied at 750 µM,the anion/cation uptake was balanced, however H⁺ excretion continuedto occur in the zone. When NO^-₃ was supplied at 5000 µM,anion uptake exceeded cation uptake but there was no OH^- excretion.Organic acid anions may be excreted by lupins to maintain theirinternal electroneutrality when anion uptake exceeds cationuptake. Rhizosphere pH would not increase unless the pK_a ofthe excreted organic anions was greater than the external pH.Copyright1993, 1999 Academic Press Lupinus angustifolius L., H⁺/OH^- excretion, nutrient uptake, cation-anion balance, vertical split root     

Failed expression of an endo-beta-1,4-glucanhydrolase (cellulase) in a non-abscinding mutant of Lupinus angustifolius cv Danja.

Cellulase expressions in a normal shedding wild-type and a non-abscinding single gene mutant of Lupinus angustifolius have been studied during ethylene treatments of leaf abscission zone explants. Of the range of different glycohydrolases investigated only the abscission cell-specific beta-1,4-glucanhydrolase (cellulase) was not produced in the non-abscinding mutant. An endo-polygalacturonase was induced equally in both wild-type and mutant and other glycohydrolases were equally up-regulated. The abscission cell-specific cellulase induced at shedding of wild-type is antigenically similar to the Phaseolus vulgaris induced leaf abscission pI 9.5 cellulase but with a higher molecular mass (50 kD compared with 48 kD) and like the bean abscission-specific cellulase that of lupin is not glycosylated. Causes of the loss of function of cellulase expression in the non-shedding mutant are discussed.     

Redistribution of Nutrient Elements from Cotyledons of Two Species of Annual Legumes During Germination and Seedling Growth

The distribution of dry matter and various mineral nutrientsbetween testa and embryo of seeds of Lupinus albus and L. angustifoliusis described It was found that lupin seeds at either end ofa pod contained less dry matter and minerals than seeds in themiddle of the fruit. The transport of dry matter, N, P, K, S,Ca, Mg, Na, Fe, Zn, Mn and Cu from cotyledons of parent seedsof both species to the seedling axis was measured from germinationto the time of cotyledon death. N, P, K and S were retrievedfrom cotyledons with over 90 per cent efficiency, dry matter,Mg, Na, Fe, Zn, Mn and with 59–90 per cent efficiency,and Ca with 26–31 per cent efficiency. There was littlechange in the efficiency of nutrient retrieval from cotyledonswhen seedlings were grown in different culture media. Both speciesshowed a linear relationship between the loss of each elementand dry matter throughout the experiment, and a similar proportioningbetween root and shoot of the amount of a specific nutrientmobilized from cotyledons of parent seeds. Lupinus albus L., Lupinus angustifolius L., lupin, transport, of dry matter and mineral nutrients     

Regulation of Stem Abscission and Callus Growth in Shoot Explants of Sweet Orange [Citrus sinensis (L.) Osbeck]

Two-node explants from Sweet Orange cv. St Ives Valencia orangeshoots produced prolific callus and formed secondary abscissionzones within internodes when cultured in vitro with abscisicacid (ABA, 5 µM) or -naphthaleneacetic acid (NAA, 5 µM).Benzyladenine (BA, 1 µm) induced callus but had littleeffect on abscission. Secondary abscission zone formation wasassociated with ABA-induced and auxin-induced ethylene formation.Treatment of explants with inhibitors of ethylene synthesis[aminoethoxyvinyl glycine (AVG), Co²⁺, PO₄^3–] preventedformation of secondary abscission zones but had variable effectson callus formation. Newly made explants contained high concentrationsof endogenous ABA (up to 6000 ng g^–1 f.wt), as measuredby GC/MS/SIM. Long-term subculture of explants (two years) inmedia containing BA (1 µm) led to a reduction in endogenousABA level (40 ng g^–1 f. wt) and to loss of capacity toform extensive callus and secondary abscission zones. Citrus sinensis (L.) Osbeck cv. St Ives Valencia, sweet orange, secondary abscission zones, in vitro, ethylene, endogenous ABA, endogenous IAA     

High pH Causes Disintegration of the Root Surface in Lupinus angustifolius L.

The effect of high pH on the morphology and anatomy of the rootsof lupin (Lupinus angustifolius L. cv. Yandee) and pea (Pisumsativum L. cv. Dundale) was examined in buffered solution. Themorphology and anatomy of lupin roots were markedly altered,and root growth was reduced by increasing solution pH from 5·2to 7·5, whereas pea roots were unaffected. In lupin roots,pH 7·5 caused disintegration of the root surface andimpaired root hair formation. Lupin roots grown at pH 7·5also had decreased cell lengths but increased cell diameterin both the epidermis and the cortex in comparison to rootsgrown at pH 5·2. High pH reduced cell volume greatlyin the epidermis, to a lesser extent in the outer cortex andnot at all in the inner cortex. It appears that in lupins, theprimary detrimental effects of growth at pH 7·5 is reducedlongitudinal growth of cells near the root surface with a consequentreduction in elongation of the cells in inner cortex.Copyright1993, 1999 Academic Press Lupinus angustifolius L., Pisum sativum L., high pH, root morphology, root anatomy     

Cellulase, Fruit Softening and Abscission in Red RaspberryRubus idaeusL. cv Glen Clova

The ripening of raspberry fruit (Rubus ideausL. cv Glen Clova)is associated with a climacteric rise in ethylene production.As the fruit pigments change from green to red there is a progressivesoftening, loss of skin strength and a breakdown of cell wallsin the mesocarp. An increase in cellulase (endo-1,4-ß-D-glucanase)in both drupelets and receptacles accompanies these changes.The localization of cellulase in the regions of the fruit associatedwith abscission zones suggest the enzyme may be involved infruit separation as well as softening. Rubus idaeusL; raspberry; fruit ripening; ethylene; abscission; cell wall breakdown; cellulase; endo-1,4-ß-D-glucanase     

Density Studies on Lupins: I. Flower Development

In an August-sown experiment the pattern of flower developmentwas followed for cv. Ultra (Lupinus albus L.) and cv. Unicrop(L. angustifolius L.) grown at low (10 plants m^–2) andhigh (93 and 83 plants m^–2, Ultra and Unicrop respectively)densities. Dry weight increase of flowers on the main-stem inflorescenceand first lateral below the main-stem were compared at differentfloral stages. Maximum flower weight was reached just priorto the open flower stage and remained constant or declined untila pod formed or abscission occurred. The time period betweenmaximum flower weight and pod formation or abscission was upto 10 days. Emergence of the inflorescence was earlier and thefirst flower of Ultra opened 10 days before Unicrop. Developmentof each terminal raceme (inflorescence) was acropetal, withpods having formed on lower flower nodes when terminal flowerswere still quite immature. Laterals forming the next generationof inflorescences grew from axillary leaf buds below an inflorescencewhile it was in full flower. Sources of competition from connectedreproductive and vegetative metabolic sinks are discussed. Lupinus spp., lupins, flower development, planting density     

Endo‐1,4‐β‐glucanase gene expression and cell wall hydrolase activities during abscission in Valencia orange

The physiological and molecular events of ethylene‐induced abscission in mature fruit calyx, laminar and floral abscission zones of cv. Valencia orange were examined. Continuous exposure of fruit explants to 5 µl 1⁻¹ ethylene for 2 to 40 h resulted in marked increases in endo‐1,4‐β‐glucanase (cellulase) and polygalacturonase (PG) activities in calyx abscission zones. Two abscission‐related cellulases and one PG were found. The major peak of cellulase activity corresponded to a pI of 8.0 and molecular weight of 51 kDa, whereas the minor cellulase peak had a pI of 5.5. The abscission polygalacturonase had a pI of 5.5. Calyx abscission zone RNA was amplified with degenerate primers based on sequence of the purified Valencia orange calyx abscission cellulase, and cloned. The two partial cellulase cDNA clones were 59% identical at the nucleotide level. Genomic Southern analysis suggested that Valencia orange contained two groups of cellulase genes. A full‐length cDNA clone from each group was isolated from a cDNA library prepared from ethylene‐induced calyx abscission zone mRNA. Both genes were expressed in ethylene‐induced calyx, laminar and floral abscission zones, but were not expressed in non‐induced abscission zones or mature leaves treated with or without ethylene, young bark or young fruit of Valencia.     

Manganese Nutrition of Lupinus spp. Especially in Relation to Developing Seeds

Patterns of transport and accumulation of manganese were studiedin Lupinus albus L. and Lupinus angustifolius L. in a wide rangeof availability levels in the rooting medium. The recently described‘split seed’ disorder, involving discolouration,splitting, and deformity of seeds, was reproduced in sand cultureusing critically low levels of manganese. The disorder was preventedby maintaining adequate manganese in the medium and its incidencein field and glasshouse was quantitatively related to the managneselevel in seed and fruit phloem sap. The use of phloem sap analysisfor early diagnosis of the disorder is suggested. High levelsof manganese in parent seed is suggested to afford protectionagainst the disorder by improving early vegetative growth ina manganese deficient situation. Direct carry-over of manganesefrom one seed generation to the next was insignificant. Manganese proved to be fully mobile in xylem but only sparinglymobile in phloem from vegetative structures to seed. It wasaccumulated in massive amounts in leaves and fruits when availabilitywas high. Seed manganese content increased 80–100 foldas the level in the rooting medium was increased from 0•1to 500 mg Mn 1^–1. L. albus was superior to L. angustifoliusin accumulating manganese in leaves and pods, and more efficientin translocating the element to its seeds. These differenceswere greatest at low or moderate manganese levels. Xylem intakeby a fruit was small relative to phloem intake when manganeseavailability was low, but became increasingly important as thesupply in the rooting medium was raised.     

Nutrient Uptake and Organic Acid Anion Metabolism in Lupins and Peas Supplied with Nitrate

Unlike many plants reported in the literature, lupins do notexcrete OH^- in amounts equivalent to the net excess of inorganicanion uptake over inorganic cation uptake. To investigate themechanisms involved in the maintenance of charge balance, nutrientuptake and organic anion accumulation of lupins and peas suppliedwith a range of NO^-₃ concentrations, were compared. Lupins absorbed less NO^-₃ than peas on a dry weight basis, whichlargely accounted for the smaller excess of anion uptake overcation uptake in lupins than in peas at the same NO^-₃ supply.When anion uptake exceeded cation uptake, peas excreted an equivalentcharge of OH^-, whereas lupins excreted much smaller amountsof OH^- than the excess of anion over cation uptake. It was calculatedthat lupins excreted significant amounts of organic anions whenanion uptake exceeded cation uptake, whereas organic anion excretionfrom peas was negligible, regardless of their NO^-₃ supply andcation-anion balance. In this study, organic anion excretion was measured from lupinroots grown in near-sterile conditions while supplied with NO^-₃at 0, 500 and 2000 µM. Although complete sterility wasnot achieved, there was close agreement between the organicanion excreted and the excess anion over cation uptake.Copyright1994, 1999 Academic Press Lupinus angustifolius L., Pisum sativum L., organic acid, nutrient uptake     

The Role of Cellulase and Polygalacturonase in Abscission of Young and Mature Shamouti Orange Fruits

During the first eight weeks after setting young citrus fruits gradually lose their ability to abscise at the abscission zone between the stem and the pedicel; in fruits older than eight weeks abscission occurs at the calyx area. The activity of cellulase and polygalacturonase in the two abscission zones was markedly increased before and during abscission, and was localized mainly in the abscission zone. Ethylene accelerated the increase in enzymic activity after an 8- to 10-h lag period; 2,4-D delayed abscission and enzymic activity when applied during the first 24 h after excision. During this period 2,4-D also partly suppressed the enhancing effect of ethylene. Early application of cyclo-heximide inhibited the formation of the enzymes and thus abscission was delayed to a certain extent. Although there are some indications that the relationship between enzymic activity and abscission is a complex one, the data presented indicate that cellulase and polygalacturonase play a significant role in abscission of citrus fruits at various developmental stages. Both enzymes act almost simultaneously and are equally controlled by ethylene and 2,4-D.     

Changes in Polygalacturonase Activity and Solubility of Polyuronides during Ethylene-stimulated Leaf Abscission in Sambucus nigra

Leaflet abscission in Sambucus nigra is precipitated by cellwall degradation which is restricted to the site of cell separation.Accompanying wall breakdown is an increase in the activity ofthe enzyme polygalacturonase (PG) (E.C. 3.2.1.15 [EC] ) and this riseis primarily confined to the abscission zone tissue. The polygalacturonasehas a pH optimum of 4·4 and has the characteristics ofan endo-acting enzyme. The elevation in enzyme activity is theresult of an increase in at least two isoforms of PG as revealedby polyacrylamide gel electrophoresis of the natured protein.Leaflet abscission in S. nigra is associated with an increasein the solubility and depolymerization of polyuronides fromthe cell wall. These observations are discussed in relationto the mechanism of cell separation during ethylene-stimulatedleaf abscission. Key words: Elder, Sambucus nigra, abscission, polygalacturonase, polyuronides, ethylene     

Effects of Ethylene and 2,4-D on the Activity of Cellulase Isoenzymes in Abscission Zones of the Developing Orange Fruit

The role of ethylene and 2,4-D in the abscission process, and the induction of cellulase isoenzymes in the abscission zones of Citrus fruit at two physiological stages of fruit development, were studied using a new staining technique for the detection of cellulase isoenzymes in polyacrylamide gels following electrophoretic separation. Four to seven isoenzymes were detected in the shoot-peduncle (zone A) and peduncle-fruit (zone C) abscission zones; at least two of them could be detected at excision time, and of these at least one could not be connected with abscission. In the young fruit, ethylene enhanced and 2,4-D delayed both abscission and the formation of several isoenzymes. In the older fruit, ethylene enhanced and 2,4-D delayed the formation of isoenzymes at a time where no abscission occurred any more in zone A. A slower but significant increase in most of the isoenzyme activity detected was also observed in abscission zone A of untreated older fruit explants after excision. These results fully agree with those reported earlier in relation to total cellulase and polygalacturonase activity (Greenberg et al., Physiol. Plant. 34: 1, 1975) tested at the same stages of fruit development. It is suggested, that the generality of the concept that a rise in hydrolytic enzymes in the abscission zone is necessarily followed by separation of the organ should be re-evaluated.     

The Role of the Stem in Phloem Loading of Minerals in Lupinus albus L. cv. Ultra

Collections of phloem sap made over a 40-day period from a varietyof locations on nodulated white lupin plants (Lupinus albusL. cv. ultra) showed considerable enrichment with K⁺ and Mg²⁺in the phloem streams destined for the shoot apices or fruitsrelative to the streams arising from the leaflets (up to 5.5times). Sodium showed enrichment in the streams destined forthe roots (up to 2.5 times) but only when present in the watersupply at a high level (3 mM). The stem, in view of its centrallocation in the transport pathway, is seen as an organ capableof redistributing minerals in the phloem independently of photosynthate. Lupinus albus L., lupin, phloem loading, magnesium, potassium, sodium, mineral elements     

Narrow-leafed Lupins with Restricted Branching

Crop phenology is one of the most important characters influencingproductivity in a given environment. Narrow-leafed lupin (Lupinusangustifolius L.) is a major grain legume crop in southern Australiawith general phenological adaptation to this Mediterranean-typeenvironment. However, it is an indeterminate crop with severalassociated limitations to productivity, such as overlappingvegetative and reproductive growth, late grain filling and sometimesexcessive vegetative growth. Here we studied two novel typesof narrow-leafed lupin with restricted branching, which mightbe useful for overcoming these problems. These restricted branchinglupins arose spontaneously within a breeding population, inthe case of ‘Tallerack’, and within a farmer's cropin the case of ‘ Hurst’ and we compared them withthe ‘Merrit’, which is widely grown and has thenormal indeterminate branching habit. The morphology and developmentof the main shoot of these genotypes were similar. However,‘Hurst’ had much larger leaves. There were alsostriking differences in the lateral branches of the restrictedbranching types; they had fewer leaves than ‘Merrit’and flowered earlier. These differences were most marked in‘ Hurst’, where the upper main stem branches werereduced to a single floret in the axil of main stem leaves,and these flowers often exhibited abnormal morphology. Copyright2000 Annals of Botany Company Lupinus angustifolius L., narrow-leafed lupin, adaptation, development, morphology, branching, leaves, mutant, plastochron, phyllochron, floral initiation, flowering.     

Effects of Diazocyclopentadiene (DACP) and Silver Thiosulphate (STS) on Ethylene Regulated Abscission of Sweet Pea Flowers (Lathyrus odoratus L.)

The ethylene production rate of cut sweet pea flower buds increased37-fold during the first 48 h of their vase life. This increasein ethylene production was accompanied by petal wilting at 72h and abscission of the buds 24 h later. Exposure of the cutspikes to the ethylene action inhibitor diazocyclopentadiene(DACP, 170 µI 1^-1) for 18 h under fluorescent lights delayedsubsequent wilting and abscission and promoted bud opening.Silver thiosulphate (0·2 mM) was more effective thanDACP, delaying wilting for longer and preventing abscissionentirely.Copyright 1995, 1999 Academic Press Ethylene, abscission, silver thiosulphate, diazocyclopentadiene, flower senescence, wilting, sweet pea, Lathyrus odoratus L     

Abscission: Response to Red and Far-Red Light

Craker, L. E., Zhao, S. Y. and Decoteau, D. R. 1987. Abscission:response to red and far-red light.—J. exp. Bot. 38: 883–888. The dose-response and time relationship of red and far-red lightin the inhibition and promotion, respectively, of dark-inducedleaf abscission was quantified using cuttings of coleus (ColeusBlumei Benth.). A continuous photon flux of approximately 15nM m^–2 s^–1 of red light was sufficient to preventleaf abscission. Abscission was promoted by exposure to a photonflux of approximately 10 nM m^–2 s^–1 of far-red lightThe inhibition of abscission by red light could be reversedby treatment with far-red and the promotion of abscission byfar-red light could be reversed by treatment with red lightThe data were consistent with a phytochrome receptor systemlocated in the leaves that controlled the presence of an abscission-inhibitingsubstance in the abscission zones. Key words: Abscission, Coleus Blumei, far-red light red light     

Using Confocal Laser Scanning Microscopy to Measure Apoplastic pH Change in Roots of Lupinus angustifolius L. in Response to High pH

A confocal laser scanning microscopy method to measure apoplasticpH in situ of the tap roots of Lupinus angustifolius L. is described.Pseudo-ratiometric imaging of cell walls of lupin tap rootswas achieved by loading living roots with a pH-sensitive dye,Cl-NERF-dextran, and a pH-insensitive dye, Texas Red-dextran,as a reference. In 5-d-old tap roots which had previously beengrown at pH 5.4, the apoplastic pH in the elongating zone was5.2 ± 0.06. The apoplastic pH in roots grown at pH 7.5increased to 5.5. The method was validated by supplying differentforms of nitrogen in the nutrient solution. With     

Density Studies on Lupins. II. Components of Seed Yield

Components of seed yield of cv. Ultra (Lupinus albus L.) andcv. Unicrop (L. angustifolius L.) were measured when grown atthree densities. The low density (10 plants m^–2) Unicropyield (34 g seed per plant) was 1.8 times that of Ultra as ithad more branches, pods and seeds per pod. Ultra seeds (310mg per seed) were heavier than Unicrop seeds (180 mg). The branchingpattern of Ultra was less dependent on plant density, henceat 93 plants m^–2 it gave a higher per plant yield (7.4vs 6.4 g) than Unicrop at lower densities (83 plants m^–2).Density had most influence on pod formation and only small effectson seeds per pod and seed weight. Yield components on the main-steminflorescence were influenced less by density than componentson branch inflorescences. Later formed, higher order generationsof inflorescences were most affected by increased inter- andintra-plant competition. Pod numbers on the main-stem were similarfor both species. Pods formed at higher flower nodes in Unicrop,but the lower flower nodes were less fertile than those in Ultra.Node position of flowers had no influence on seed set in main-stemUnicrop pods, but pods from higher nodes in Ultra formed fewerseeds. Seed weights in Unicrop were similar among main-stemnodes but in Ultra seed weights tended to increase at highernodes. Lupinus spp, lupins, seed yield, planting density