OXYNOE AZUROPUNCTATA, N. SP., A NEW SACOGLOSSAN FROM THE FLORIDA KEYS (MOLLUSCA: OPISTHOBRANCHIA)

A new species of Oxynoe from the Florida Keys is described.It differs from the other Caribbean Oxynoe, O. antillarum Mörch,1863, in color, number and size of papillae, size increase ofradular teeth, diet, and type of development. The type of developmentis extended lecithotrophic. The larvae hatch about 4 weeks afterthe eggs have been deposited, and metamorphosis is completedafter about one additional week. Oxynoe azuropunctata n.sp.feeds on Caulerpa paspaloides, C. cupressoides and C. sertularioides.Details of the feeding process in Oxynoe are described for thefirst time. The junction of the oral tube and the pharynx isequipped with a complex muscular buccal apparatus, part of whichis everted during feeding. The protrusible parts of the buccalapparatus completely surround part of an algal filament, whichis then slit with the radula, and the cell sap is sucked out.The protrusible parts are retracted after the filament has beenreleased. ^*Present address: Ellebjergvej 22, 1 t.h., DK-2450 CopenhagenSV, Denmark (Received 13 November 1979;     

STUDIES ON NITELLA HAVING ARTIFICIAL CELL SAP I. REPLACEMENT OF THE CELL SAP WITH ARTIFICIAL SOLUTIONS

A method for replacing the cell sap of Nitella with an artificialsolution was introduced. The technique, which is a modificationof KAMIYA and KURODA'S (1, 2), is applicable not only for isotonicbut also for hypertonic or hypotonic solutions. Photometricdeterminations of K⁺, Na⁺, Ca⁺⁺ and Cl^– proved that thereplacement of the cell sap with the present method is satisfactory.The internodal cell of Nitella, whose cell sap was replacedwith an isotonic solution with a simple composition such asa mixture of KCl, NaCl and CaCl₂, can be kept living at leastfor several days, sometimes even for more than one month. (Received September 6, 1963; )     

Inhibition of Malate Synthesis by Vanadate: Implications for the Cytosolic Alkalinization Induced by Vanadate Concentrations Partially Inhibiting the Plasmalemma H+ Pump

The effects of Na-orthovanadate, at concentrations only partiallyinhibiting net H⁺ extrusion, were determined on vacuolar andcytosolic pH by the weak base and weak acid distribution atequilibrium. Treatment with vanadate induces in Elodea densaleaves and in Arabidopsis thaliana seedlings a moderate acidificationof both cell sap and vacuole. Conversely, it induces an alkalinizationof cytosol, this effect being in apparent contrast with a conditionof reduced activity of the H⁺-transporting plasmalemma ATPase,which should be associated with a cytosolic acidification. InArabidopsis seedlings treated with vanadate, the increase inpH of both cytosol and external medium is associated with adecrease in cell sap buffer capacity, more evident for highervanadate concentrations, and particularly marked in the pH rangebetween 3·5 and 5·5. In these conditions, themalate content is strongly reduced, its decrease almost completelyaccounting for the decrease in cell sap buffer capacity. Anin vitro analysis of the vanadate effect on phosphoenolpyruvatecarboxylase indicates that the decrease in malate content seemssubstantially due to an inhibiting effect of vanadate on thisenzyme. These results stress that the in vivo use of vanadateas an inhibitor of the plasmalemma H⁺-ATPase must be taken withcaution; in particular, for studying the correlations betweenchanges in net H⁺ extrusion and changes in cytosolic pH andrelated processes. Key words: Vanadate, malate, cytosolic pH, Elodea densa, Arabidopsis thaliana     

A 15N Method to Quantify the Sources of Nitrogen for New Growth of Non-legumes, Examined using Kiwifruit

A new method is described for estimating the contributions ofnitrate-N, newly-reduced nitrate-N and remobilization of storednitrogen (N) for new growth of non-leguminous plants. Xylemsap is extracted from ¹⁵N-labelled and unlabelled plants. Nitrate-Nand amino-N in sap are separated using cation-exchange resinand analysed for ¹⁵N. Remobilization of stored N is estimatedusing [I-(¹⁵N enrichment of amino-N/¹⁵N enrichment of nitrate-N)]x amino-N. Newly-reduced nitrate-N is estimated by difference.Field-application of this method is described. Actinidia deliciosa, kiwifruit, nitrogen, ¹⁵N, remobilization, xylem sap, nitrate reduction     

IONIC COMPOSITION AND ELECTRIC RESPONSE OF LAMPROTHAMNIUM SUCCINCTUM

The concentrations of K⁺, Na⁺ and Cl^– in the cytoplasmof Lamprothamnium succinctum, a brackish water Characeae, areabout 137, 47 and 86 mM respectively. The concentration of K⁺in the cytoplasm is of the same order as that in the cell sap,while the concentrations of Na⁺ and Cl^– are much lowerthan those in the cell sap (i.e., 140 mM Na⁺, 370 mM Cl^–).In the brackish water, in which the plant grows, the internodesis never excitable electrically. However, it acquires excitabilitywhen it is kept in a mannitol-pond water. The action potentialthus elicited is accompanied by a temporary cessation or slowdown of the cytoplasmic streaming. ¹This work was supported by Research Grants from the Ministryof Education of Japan     

A Reanalysis of Particle Motion in Sieve Tubes of Heracleum

From an analysis of the Brownian motion of particles in sieveelements of Heracleum mantegazzianum and Heracleum sphondylium,Barclay and Johnson have suggested that the in situ viscosityof sieve tube sap is four to six times higher than has previouslybeen assumed. In particular, they obtained a value for the sapviscosity of about 10^–2 Pa s, which compares with a valueof 2 x 10^–3 Pa s for a 20 per cent (w/v) sucrose solution.The present paper describes a reanalysis of their data. It isargued that Barclay & Johnson underestimated the Brownianmotion of sieve element particles and so overestimated the sapviscosity. An exact correction was not possible, but it is concludedthat the in situ viscosity of Heracleum sieve tube sap mustbe less than 3 x 10^–3 Pa s, which corresponds to a sucroseconcentration of less than 29 per cent. Hence it may not beunreasonable to suppose that the viscosity of sieve tube sapis determined primarily by the concentration of sucrose, ashas been assumed in theoretical analyses of the Munch hypothesis.It is also concluded that the sieve tubes studied by Barclayand Johnson were not functional, in the sense that they didnot exhibit an axial bulk flow of sap. Heracleum, sieve tubes, Brownian motion, viscosity, Munch hypothesis     

Lysosomal nature of plant vacuoles II. Acid hydrolases in the central vacuole of internodal cells of Charophyta

Contents of the central vacuole (cell sap) were separated frominternodal cells of Nitella and Chara. Most of the acid phosphataseand carboxypeptidase, which are marker enzymes for lysosomes,were detected in the separated cell sap. In contrast, most ofthe catalase, cytochrome oxidase, NADPH₂-cytochrome c reductase,and glucose-6-phosphate dehydrogenase were detected in partsof the cell other than the cell sap: which shows there is relativelylittle contamination of the cytoplasm in the separated cellsap. Enzymatic properties of the carboxypeptidase in Nitellaaxilliformis were similar to those of carboxypeptidases in higherplants and those of cathepsin A in animal lysosomes. These resultsindicate that the central vacuole of Charophyta has some propertiesof the lysosome. ¹ Preliminary results of this paper were given in reference(5). (Received February 3, 1975; )     

Feeding behaviour of the rotifer Ascomorpha ovalis: functional response, handling time and exploitation of individual Ceratium cells

The planktonic rotifer Ascomorpha ovalis feeds on large dinoflagellates(e.g. Ceratium sp., Peridinium sp.) and is able to extract theircell contents by means of its virgate mastax. This paper presentsthe results of experiments on the feeding behaviour of laboratory-culturedAscomorpha with Cerarium furcoides as food algae. Ascomorphaare three times larger than their prey Ceratium (by volume),but with regard to total length, their prey was even 20% larger.Ascomorpha showed a hyperbolic functional response curve witha plateau of the feeding rate at 8 Ceratium cells animal^–1dar^–1 when concentrations of Ceratium were >100 cellsml^–1. The mean handling time (time for capturing and extractingone Ceratium cell) was 3 min. The shape of the functional responsewas better described by a curvilinear model than by a rectilinearmodel. However, handling times cannot be responsible for this,since they were too short to set limits on ingestion rates.At low food concentrations, encounter rates with prey seemedto limit the feeding rates of Ascomorpha, whereas at mediumto high food concentrations, Satiation effects (lower attackrates) seemed to set limits on the feeding rates. Ascomorphashowed a significant decrease in the exploitation of singleCeratium cells at high prey concentrations. This decrease couldbe explained by a saturation effect in which the partly filledguts of Ascomorpha did not permit the total extraction of thecontents of a Ceratium cell.     

Haustorium-related Uptake and Metabolism of Host Xylem Solutes by the Root Hemiparasitic ShrubSantalum acuminatum(R. Br.) A. DC. (Santalaceae)

Solute composition of root xylem sap of common native hostsof quandong (Santalum acuminatum) was compared with that ofcorresponding xylem sap and ethanolic extracts of endophytictissues of haustoria of the hemiparasite. Each host transporteda characteristic set of organic nitrogenous solutes, but littleor no nitrate, and the data indicated only limited direct flowof amino compounds between xylem streams of hosts and parasite.Proline predominated in the haustorium and xylem ofSantalum,but was at negligible levels in the xylem of most hosts. Sucrose,fructose, glucose, malate and citrate were at high levels inall saps, and fructose especially prominent inSantalum. Chloride,sulphate and phosphate were the principal inorganic anions ofthe xylem. Based on C:N ratios of xylem and dry matter ofSantalumandassuming a 70% or more dependence on the host for N, it wasestimated thatSantalumwould gain approximately one third ofits C requirement for dry matter production heterotrophicallyfrom the xylem of its hosts. Infiltration of xylem of haustoria-bearingroot segments of a major host (Acacia rostellifera) with a rangeof¹⁵N labelled substrates resulted in 40–80% of the¹⁵Nof endophytes of the attached haustoria being received as proline.Nitrate reductase activity was induced in haustoria followinghost xylem feeding of nitrate. The study concludes that haustoriaofSantalumact as a major site of synthesis and export of prolineand might therefore play an important role in osmotic adjustmentof the parasite and its related acquisition of water from hosts. Root hemiparasite; Santalum acuminatum; ¹⁵N labelled substrates; xylem transport; proline; osmoregulation     

Feeding, growth and food conversion of the marine cladoceran Penilia avirostris

Ingestion and clearance rates, feeding behaviors and life historyvariables of the marine cladoceran Penilia avirostris were evaluatedover a range of food concentrations encountered in nature (0.01–3.0mm³ 1^–1 of Isochrysis galbana). Ingestion rates increasedand clearance rates decreased with increasing food concentration.No maximum feeding thresholds were observed over the range ofalgal concentrations offered. Weight-specific ingestion ratesdecreased with increasing body weight. Feeding behaviors suchas mandibular activity, abdominal claw rejections of cloggedfeeding structures and feeding appendage activity decreasedat a food level of 0.3 mm³ l^–1 of l. galbana. Peniliaavirostris had very poor survivorship at extremely low (0.01mm³ l^–1 and high (3.00 mm³ l^–1) food levels. Mortalitywas hardly affected at food levels of 0.03–1.0 mm³ l^–1Reproduction did not occur at food levels of     

Effects of prey motility and concentration on feeding in Acartia tonsa and Temora longicornis: the importance of feeding modes

Feeding experiments were conducted with the ambush-feeding copepodAcartia tonsa and the feeding-current-generating copepod Temoralongicornis. The copepods were offered a mixed diet of the dinoflagellateHeterocapsa triquetra and the ciliate Balanion comatum of similarcell size. The dinoflagellate was offered at a constant concentrationof 10–15 cells mL^–1, whereas the ciliate was offeredat a variety of concentrations, ranging from 7 to 57 cells mL^–1.Copepods with different feeding modes possess different mechanismsfor prey detection, suggesting that the two copepods would responddifferently to the two prey types. Both copepods had significantlyhigher clearance rates on the highly motile ciliate than onthe less motile dinoflagellate. In encounters between A. tonsaand its prey, we argue that this is due to the higher hydromechanicalsignal generated by the ciliate. The advection feeding copepodT. longicornis fed on the two prey according to their relativeconcentrations; in this case, we suggest that although B. comatumis capable of detecting feeding-current-generating predators,the feeding current velocity generated by T. longicornis isgreater than the escape velocity of this ciliate.     

Stomatal response to abscisic acid fed into the xylem of intact Helianthus annuus (L.) plants

The effect of abscisic acid on stomatal apertures of sunflower(Helianthus annuus (L.)) was investigated with a new methodfor feeding solutions into an attached leaf of an intact plant.Xylem sap was sampled with a Passioura-type pressure chamber.Then it was modified in its composition and fed back into amature leaf of the plant from which it had been collected beforethe experiment. Simultaneously, unmodified xylem sap was fedinto a control leaf at the same internode. The use of the Passioura-typepressure chamber during feeding, prevented embolisms and ensuredminimum dilution of the feeding solution. The effect of feedingwas measured by two gas exchange systems, located at the treatmentand at the control leaf. During the feeding experiments up to84% of the water volume transpired by the leaf was substitutedby the supplied feeding sap. When feeding xylem sap, to which2.5 mmol m^–3 ABA (physiological range) was added, leafconductance decreased to a similar value as in drought experiments.A log-linear relationship between the fed ABA-concentrationand leaf conductance was observed. Low stomatal con-ductancewas dependent on a continuous supply of ABA to the leaf. Whentotal ABA-influx into the leaf was large, either due to long-termfeeding of low concentrations or short-term feeding of highconcentrations (i) recovery after feeding started later and(ii) the rate of recovery was decreased. Therefore, stomatalresponses after short-term and long-term ABA-feeding were dependenton the loading of ABA into the leaf and not only on ABA-concentrations.The effectiveness of fed ABA was also dependent on the lightintensity at the fed leaf. Key words: Abscisic acid, feeding method, stomata, gas exchange, Helianthus annuus     

Characterization of Cd translocation and identification of the Cd form in xylem sap of the Cd-hyperaccumulator Arabidopsis halleri

Arabidopsis halleri is a Cd hyperaccumulator; however, the mechanismsinvolved in the root to shoot translocation of Cd are not wellunderstood. In this study, we characterized Cd transfer fromthe root medium to xylem in this species. Arabidopsis halleriaccumulated 1,500 mg kg^–1 Cd in the shoot without growthinhibition. A time-course experiment showed that the releaseof Cd into the xylem was very rapid; by 2 h exposure to Cd,Cd concentration in the xylem sap was 5-fold higher than thatin the external solution. The concentration of Cd in the xylemsap increased linearly with increasing Cd concentration in theexternal solution. Cd transfer to the xylem was completely inhibitedby the metabolic inhibitor carbonyl cyanide 3-chlorophenylhydrazone(CCCP). Cd concentration in the xylem sap was decreased by increasingthe concentration of external Zn, but enhanced by Fe deficiencytreatment. Analysis with ¹¹³Cd-nuclear magnetic resonance (NMR)showed that the chemical shift of ¹¹³Cd in the xylem sap wasthe same as that of Cd(NO₃)₂. Metal speciation with Geochem-PCalso showed that Cd occurred mainly in the free ionic form inthe xylem sap. These results suggest that Cd transfer from theroot medium to the xylem in A. halleri is an energy-dependentprocess that is partly shared with Zn and/or Fe transport. Furthermore,Cd is translocated from roots to shoots in inorganic forms.     

Effect of soil compaction on barley (Hordeum vulgare L.) growth: II Are increased xylem sap ABA concentrations involved in maintaining leaf expansion in compacted soils?

The abscisic acid (ABA)-deficient mutant of barley, Az34, exhibiteda much reduced rate of leaf expansion at a bulk density of 1.6g cm^–3 as compared to the isogenic wild-type variety,Steptoe. Az34 had a consistently lower xylem sap ABA concentrationat 7 d and 14 d after emergence (DAE). The xylem sap data suggestthat ABA present at Steptoe concentrations may have a directrole in maintaining leaf expansion at the sub-critical bulkdensity (1.6 g cm^–3 To test this hypothesis, additionof synthetic ABA either to the rooting environment (100 nM)or directly to the xylem sap (5 pg µl^–1 to reproducethe xylem sap ABA concentrations found in Steptoe, increasedleaf expansion in Az34 to the wild-type level. Furthermore,feeding Steptoe xylem sap to Az34 produced similar effects.These experiments provide direct evidence of a positive rolefor ABA as a root-to-shoot signal which assists in maintainingleaf growth in plants experiencing subcritical levels of compactionstress. Key words: ABA-deficient mutant, leaf expansion, xylem sap, ABA     

MONOPHASIC AND DIPHASIC DIGESTIVE CYCLES IN VENERUPIS DECUSSATA AND CHLAMYS VARIA

Feeding and digestive cycles in Chlamys varia and Venerupisdecussata are identified and shown to relate to a tidal cycle.The processes of digestion within the diverticula of the twospecies are very different. The tubules of Venerupis are synchronizedand exhibit a monophasic cycle completed within a twelve-hourperiod, facilitating feeding at the next cycle of the tide.The digestive process of Chlamys requires twenty-four hoursand to accommodate feeding at each twelve-hour tidal cycle thetubules within the diverticula exhibit two different digestivephases simultaneously so that a diaphasic cycle is apparent.The pH of the mantle cavity and regions of the digestive tractand the formation and dissolution of the crystalline style areshown to be related to the tidal cycle. ^*Department of Brewing and Biological Sciences, Heriot-WattUniversity, Chambers Street, Edinburgh, EH1 1HX. (Received 22 March 1978;     

Feeding, prey selection and prey encounter mechanisms in the heterotrophic dinoflagellate Noctiluca scintillans

The heterotrophic dinoflagellate Noctiluca scintillans has anegligible swimming ability and feeds predominantly on immobileprey. How, then, does it encounter prey? Noctiluca scintillansis positively buoyant and, therefore, we hypothesized that itintercepts prey particles during ascent and/or that microscaleshear brings it into contact with prey. Noctiluca scintillanshas a specific carbon content 1–2 orders of magnitudeless than that typical for protists and, thus, an inflated volume.It also has a density slightly less than that of the ambientwater and therefore ascends at high velocities (-1 m h^–1).In stagnant water, clearance rates of latex spheres (5–80µm) increased approximately with prey particle size squared.This scaling is consistent with N.scintillans being an interceptionfeeder. However, absolute clearance rates were substantiallylower than those predicted by modeling N.scintillans both asa spherical and as a cylindrical collector. The latter modelassumes that prey particles are collected on the string of mucusthat may form at the tip of the tentacle. Feeding, growth andprey selection experiments all demonstrated that diatoms arecleared at substantially higher rates than latex beads and otherphytoplankters, particularly dinoflagellates. We propose thatdiatoms stick more efficiently than latex beads to the mucusof N.scintillans and that dinoflagellates reduce fatal contactbehaviorally. We conclude that N.scintillans is an interceptionfeeder and that the high ascent velocity accounts for encounterswith prey. However, the flow field around the cell-mucus complexis too complicated to be described accurately by simple geometricmodels. Fluid shear (0.7–1.8 s^–1 had a negativeimpact on feeding rates, which were much less than predictedby models. Noctiluca scintillans can survive starvation forlong periods (>3 weeks), it can grow at low concentrationsof prey (-15 µg C l^–1), but growth saturates onlyat very high prey concentrations of 500–1000 µgC l^–1 or more. We demonstrate how the functional biologyof N.scintillans is consistent with its spatial and seasonaldistribution, which is characterized by persistence in the plankton,blooms in association with high concentrations of diatoms, andsurface accumulation during quiescent periods or exponentialdecline in abundance with depth during periods of turbulentmixing.     

FEEDING ACTIVITY OF LIMAX VALENTIANUS FERUSSAC: NOCTURNAL RHYTHM AND ALIMENTARY COMPETITION

Feeding activity of the slug Limax valentianus Férus-sacthroughout a 24-hour period began with the decrease in light.Feeding began 41 minutes after the start of locomotor activityand showed a maximum level in the beginning of the night L.valentianus had two to five meals each night, each meal lasting8.5 to 14 minutes. The overcrowding of the slugs induced a shorterduration of individual feeding periods and acts of aggressionduring meals. ^*Present address: Centre de Neurochimie, Laboratoire de neurobiologiemolécuiaire des interactions cellulaires. 5 rue BlaisePascal, 67084 Strasbourg, France (Received 7 July 1996; accepted 21 April 1997)     

The Composition of Phloem Exudate and Xylem Sap from Tree Tobacco (Nicotiana glauca Grah.)

The composition of xylem sap and exudate from stem incisionsof Nicotiana glauca Grah. was compared in detail. Exudationfrom stem incisions occurred over a 5 min period in certainplants, enabling collection of 5–30 µl of sap. Therate of exudation showed an exponential decline. Exudate hada high dry matter content (170–196 mg ml^–1) andhigh sugar (sucrose) levels. Xylem sap had a low pH (5.8) andexudate a pH of 7.9. Glutamine dominated the amino compoundsin xylem sap and exudate, and K⁺ was the major cation. Totalamino compounds in stem exudate reached 10.8 mg ml^–1 whereasxylem sap contained much lower levels (0.28 mg ml^–1).All mineral elements and amino compounds with the exceptionof calcium were more concentrated in stem exudate than in xylemsap. Sucrose was labelled heavily in stem exudate following pulsingof an adjacent leaf with ¹⁴CO₂. A concentration gradient ofsugar (2.1 bar m^–1) was recorded for stems. Levels ofsucrose, amino compounds and K⁺ ions in stem exudate showeda diurnal periodicity. Each commodity reached maximum concentrationat or near noon and minimum concentration about dawn. The evidencesuggests that exudate from stem incisions of N. glauca is arepresentative sample of solutes translocated in the phloem. Nicotiana glauca Grah., phloem sap, xylem sap, sucrose, amino compounds, mineral ions     

Transient response of sap flow to wind speed

Transient responses of sap flow to step changes in wind speedwere experimentally investigated in a wind tunnel. A Granier-typesap flow sensor was calibrated and tested in a cylindrical tubefor analysis of its transient time response. Then the sensorwas used to measure the transient response of a well-wateredPachira macrocarpa plant to wind speed variations. The transientresponse of sap flow was described using the resistance–capacitancemodel. The steady sap flow rate increased as the wind speedincreased at low wind speeds. Once the wind speed exceeded 8.0m s^–1, the steady sap flow rate did not increase further.The transpiration rate, measured gravimetrically, showed a similartrend. The response of nocturnal sap flow to wind speed variationwas also measured and compared with the results in the daytime.Under the same wind speed, the steady sap flow rate was smallerthan that in the daytime, indicating differences between diurnaland nocturnal hydraulic function, and incomplete stomatal closureat night. In addition, it was found that the temporal responseof the Granier sensor is fast enough to resolve the transientbehaviour of water flux in plant tissue. Key words: Nocturnal transpiration, sap flow, transient response, wind speed Received 31 July 2008; Revised 6 October 2008 Accepted 20 October 2008     

The Cellular Pathway of Short-distance Transfer of Photosynthates and Potassium in the Elongating Stem ofPhaseolus vulgarisL. A Physiological Assessment

Plasmolytic disruption of plasmodesmata interconnecting metaphloemsieve element-companion cell complexes with small and largephloem parenchyma cells in the elongating region of internode2 ofPhaseolus vulgarisL. seedlings did not affect accumulationof phloem-imported¹⁴C-photosynthates and⁸⁶rubidium. The membrane-impermeantdye, 5(6) carboxyfluorescein, loaded into leaf phloem as themembrane-permeant diacetate ester, was found not to move radiallyout of the importing sieve elements in the internode elongationregion. In contrast, the apoplasmic tracer, Calcuofluor White,rapidly moved laterally throughout all tissues of the elongationzone. Hexoses, sucrose and potassium were identified as themain osmotica in internode apoplasmic sap. Label asymmetry in[¹⁴C](fructosyl)sucrose was retained on accumulation by excisedstem segments. Uptake of [¹⁴C]sucrose and⁸⁶rubidium by stemsegments exhibited saturation kinetics. Sucrose uptake was inhibitedby the slowly penetrating sulphydryl reagent, para-chloromercuribenzenesulphonicacid.In vitrorates of sucrose uptake, at apoplasmic concentrations,corresponded to its predictedin vivorate of delivery to thestem ground tissues from mature sieve elements when respiratorylosses were assumed to be confined to the stem phloem. For potassium,the total delivery rate could be accounted for by itsin vitrorateof uptake. Overall, it was concluded that radial transport,in the elongation zone of internode 2 ofPhaseolus vulgarisL.seedlings, follows an apoplasmic route from mature sieve elementsto stem ground tissues.Copyright 1998 Annals of Botany Company PhaseoluLes vulgaris, apoplasm, elongating stem, French bean, photosynthates, potassium, radial transfer, symplasm.