Ecophysiological Aspects of the Woody Root HemiparasiteSantalum acuminatum(R. Br.) A. DC and its Common Hosts in South Western Australia

The water and nitrogen relationships of the xylem-tapping roothemiparasite, quandong (Santalum acuminatum) and its principalhosts were examined at a series of sites in native coastal heathlandsof south west Australia. Assessments based on densities of above-groundbiomass, ground cover and frequencies of haustoria on host rootsindicated that woody N₂fixers (legumes andAllocasuarina) wereprincipal hosts ofSantalum. ¹⁵N values for shoot dry matterof component species suggested these N₂fixers were strongly(70% or more) dependent on atmospheric N and thatSantalumderivedN principally from these species. Structural studies of haustoriashowed the interface with host xylem to be almost entirely comprisedof parenchymatous tissue. No luminal continuities were observedbetween xylem conducting elements of the partners. Formationand functional life of haustoria were closely coordinated withseasonal growth of hosts, with some haustoria surviving summerand overlapping functionally with new ones establishing in thenext autumn. Transpiration and photosynthetic rates of the parasitewere consistently less and water use efficiencies very similarto those of the principal hostAcacia rostellifera. ¹³C valuesof foliage ofSantalumand this host were similar, but large variationsin ¹³C values for above-ground dry matter of parasite and hostsbetween study sites prevented evaluations of water stress orwater-use-efficiency based on carbon isotope discrimination.Specific hydraulic conductivities of roots ofSantalumwere consistentlylower than those ofAcacia, a finding consistent with more conservativewater use by the parasite than the host. Santalum acuminatum; root hemiparasite; ¹⁵N discrimination; water relations; haustorial structure; root conductivity     

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     

Photosynthesis and Transpiration in Rice as Influenced by Soil Moisture and Air Humidity

The paper describes the effect of soil moisture content andair humidity on CO₂ exchange (P_N), CO₂ diffusion resistance(Cr) and transpiration (E) in four varieties of japonica rice(Oryza sativa L.). A decrease in soil moisture content reducedthe rate of photosynthesis to a varying degree in the varieties.Reduction in photosynthesis was attributed to increase in Cr.The effect of low soil moisture on photosynthesis and CO₂ diffusionwas further intensified by decrease in air humidity. By maintaininga high humidity in the air around the leaves however, the effectof soil moisture deficiency was reduced considerably, exceptin Rikuto Norin 21 which was very sensitive to soil-moisturedeficiency alone. Dryness of the air enhanced the transpirationrate, although the increase was relatively less in the plantsfacing a simultaneous water crisis at the root surface. In plantsgrowing under flooded conditions, a decrease in air humiditycaused a slight depression in P_N despite the simultaneous decreasein Cr. Oryza sativa L., rice, photosynthesis, transpiration, diffusion resistance, soil moisture, air humidity     

Growth of Actinorhizal Plants as Influenced by the Form of N with Special Reference to Myrica gale and Alnus incana

Growth of two actinorhizal species was studied in relation tothe form of N supply in water culture. Non-nodulated bog myrtle(Myrica gale) and grey alder (Alnus incana) were grown withNH₄⁺, NH₄NO₃^– or NO₃^– (4 mol m^–3 N). A nodulatedseries of bog myrtle was also cultivated in N-free medium. Relative growth rate (RGR), utilization rate of N, and shoot/rootratio were highest for the two species with the N completelysupplied as NH₄⁺. In both species, nitrate was largely reducedin the roots and the presence of NO₃^– in combined-N supplyalways affected the RGR and N utilization rate negatively. BothN₂ fixation and complete NO₃^– nutrition represented conditionsof relative N-deficiency resulting in relatively low tissue-Nconcentrations and a greater allocation of dry mass to the roots.The physiological N status of nodulated M. gale plants was comparativelygood, as indicated by a normal nodule weight ratio and a relativelyhigh N₂-fixing rate per unit nodule mass. However, whole-plantN₂-fixing capacity remained relatively low in comparison withacquisition rates of N in combined-N plants. The anion charge from the nitrate reduction was largely directlyexcreted as an OH^– efflux. H ⁺ /N ratios generally agreedwith the theory. In comparison with NH₄⁺ nutrition, carboxylateconcentrations were higher in N₂-fixing M. gale plants and theH ⁺ /N ratio in nodulated plants was less than unity below thevalue for ammonium plants as previously found for other actinorhizalspecies. Therefore, NH₄⁺ should be an energetically more attractiveN source for actinorhizal plants than N₂. The results agree with commonly accepted views on energeticsof N uptake and assimilation in higher plants and support theconcept of a basically similar physiological behaviour betweennon-legumes and legumes. Key words: Actinorhizal symbioses, ammonium, H⁺/OH^– efflux, nitrate, N₂ fixation, NRA     

Influence of High CO2Partial Pressure on Nitrogen Use Efficiency of the C4Grasses Panicum coloratum and Cenchrus ciliaris

Australia's tropical grasslands are dominated by C₄grasses,characterized by their unique biochemistry and anatomy. Twonaturalized C₄grasses (Panicum coloratum and Cenchrus ciliaris)were used to investigate whether high CO₂partial pressure [p(CO₂)] influences photosynthetic nitrogen use efficiency andplant nitrogen use efficiency (PNUE and NUE respectively). Plantswere grown for 30 d with four levels of N at p(CO₂) of 38 or86 Pa. PNUE was calculated from leaf CO₂assimilation rates (A)and leaf N concentrations, and NUE from total leaf N contentand plant dry mass. At each p(CO₂), PNUE and NUE were greaterfor C. ciliaris than for P. coloratum due to higher A and drymass combined with lower leaf N concentrations. Elevatedp (CO₂)increased PNUE of C. ciliaris only. This effect was due to lowerleaf N concentrations (area basis). At high p(CO₂), NUE of C.ciliaris was also greater. This resulted from a 1.6-fold stimulationof dry mass by high p(CO₂). Although dry mass of P. coloratumwas increased 1.2-fold by elevated p(CO₂), its NUE was unaffected.Leaf transpiration rates were halved at elevated p(CO₂), andwe suggest that this factor plays a major role in the growthresponse of C₄grasses to high p(CO₂). Copyright 2001 Annalsof Botany Company Panicum coloratum, Cenchrus ciliaris, nitrogen use efficiency, elevated CO₂, leaf N concentration, growth, photosynthesis     

Effects of the fungal endophyte, Neotyphodium lolii, on net photosynthesis and growth rates of perennial ryegrass (Lolium perenne) are independent of In Planta endophyte concentration

• Background and Aims Neotyphodium lolii is a fungal endophyteof perennial ryegrass (Lolium perenne), improving grass fitnessthrough production of bioactive alkaloids. Neotyphodium speciescan also affect growth and physiology of their host grasses(family Poaceae, sub-family Pooideae), but little is known aboutthe mechanisms. This study examined the effect of N. lolii onnet photosynthesis (P_n) and growth rates in ryegrass genotypesdiffering in endophyte concentration in all leaf tissues. • Methods Plants from two ryegrass genotypes, Nui D andNui UIV, infected with N. lolii (E+) differing approx. 2-foldin endophyte concentration or uninfected clones thereof (E–)were grown in a controlled environment. For each genotype xendophyte treatment, plant growth rates were assessed as tilleringand leaf extension rates, and the light response of P_n, darkrespiration and transpiration measured in leaves of young (30–45d old) and old (>90 d old) plants with a single-chamber openinfrared gas-exchange system. • Key Results Neotyphodium lolii affected CO₂-limited ratesof P_n, which were approx. 17 % lower in E+ than E– plants(P < 0·05) in the young plants. Apparent photon yieldand dark respiration were unaffected by the endophyte (P >0·05). Neotyphodium lolii also decreased transpiration(P < 0·05), but only in complete darkness. There wereno endophyte effects on P_n in the old plants (P > 0·05).E+ plants grew faster immediately after replanting (P < 0·05),but had approx. 10 % lower growth rates during mid-log growth(P < 0·05) than E– plants, but there was noeffect on final plant biomass (P > 0·05). The endophyteeffects on P_n and growth tended to be more pronounced in NuiUIV, despite having a lower endophyte concentration than NuiD. • Conclusions Neotyphodium lolii affects CO₂ fixation,but not light interception and photochemistry of P_n. The impactof N. lolii on plant growth and photosynthesis is independentof endophyte concentration in the plant, suggesting that theendophyte mycelium is not simply an energy drain to the plant.However, the endophyte effects on P_n and plant growth are stronglydependent on the plant growth phase.     

Host specific variation in photosynthesis of an obligate xylem‐tapping mistletoe Dendrophthoe curvata in a Bornean heath forest

Most mistletoe–host ecophysiological studies have paid attention to the influence of parasites on host performance. This paper explored the impact of varying hosts on the photosynthesis of a single mistletoe species. Here, we studied an obligate xylem‐tapping tropical mistletoe (Dendrophthoe curvata (Blume) Miquel) parasitizing four different hosts (Acacia auriculiformis A. Cunn. Ex Benth, Andira inermis (W. Wright) DC., Mangifera indica L. and Vitex pinnata L.) in a homo geneous tropical heath forest patch in Brunei Darussalam. We compared photosynthetic capacity and photosynthesis‐related characteristics of the mistletoe on four different hosts to evaluate the overall impact of hosts on the parasite. Results showed that the mistletoe–host patterns of CO₂ assimilation rates, transpiration rates and water use efficiency varied significantly based on the host. In the D. curvata–Vitex pinnata association, the mistletoe exhibited significantly lower CO₂ assimilation rates but showed no significant variations in transpiration rates and water use efficiency when compared to the host. In D. curvata–Andira inermis and D. curvata–Mangifera indica associations, the mistletoe showed significantly higher photosynthetic rates than the hosts, whereas in the D. curvata–Acacia auriculiformis association, there was no significant difference in photosynthetic rates between the counterparts. Host specificity also significantly influenced some mistletoe photosynthetic parameters such as light saturated photosynthesis, specific leaf area, leaf chlorophyll content, CO₂ assimilation rates, stomatal conductance, transpiration rates and water use efficiency. Different tree hosts intrinsically offer different resources to their obligate mistletoe parasites based on their physiology and environmental parameters. We argue that host‐specific responses have driven these intra‐specific variations in mistletoe physiology. This study provides background for future investigation on potential host‐regulated mechanisms that drive functional changes in host‐dependent mistletoes.     

Influence of nitrogen on growth and photosynthesis of a C3 cereal, Oryza sativa, infected with the root hemiparasite Striga hermonthica

Striga hermonthica is a root hemiparasitic angiosperm nativeto the African semi-arid tropics. It is a major weed of C₄ cerealsbut locally it is also an important weed of the C₃ plant, rice[Oryza sativa). Infected rice plants produced 17% and 42% ofthe total biomass of uninfected plants when grown at two differentammonium nitrate concentrations, 1 and 3 mol m^–3, respectively.S. hermonthica prevented grain production at both concentrationsof nitrogen. At the lower concentration no heads were produced.At the higher concentration head weight was only 6% of uninfectedcontrols. S. hermonthica also altered the partitioning of drymatter between plant parts, such that shoot growth was reducedto a greater extent than root growth. As a consequence the root-to-shootratio of infected plants was approximately five times greaterthan that of uninfected control plants. Light saturated ratesof photosynthesis In infected plants were 56% and 70% of thoseof uninfected controls, at low and high nitrogen, respectively.Infection also led to lower values of stomatal conductance althoughthe substom-atal CO₂ concentration was unaffected. Analysisof the response of photosynthesis to substomatal CO₂ concentration(A/C_I curves) demonstrated that lower rates of photosynthesiscould not be solely attributed to lower stomatal conductances.Lower initial slopes and asymptotic rates suggest that bothcarboxylation and processes controlling regeneration of ribulose-1,5-bisphosphate are reduced by infection. The data are discussedwith respect to the influence of S. hermonthica on the growthand photosynthesis of C₄ hosts, where in contrast to the situationwith rice, nitrogen feeding results in a marked alleviationof the effects of the parasite on the host. Key words: Rice, Striga, growth, photosynthesis, nitrogen     

Nitrogen Fixation in the Phyllosphere of Tropical Plants: Occurrence of Phyllosphere Nitrogen-Fixing Micro-organisms in Eastern India and their Utility for the Growth and Nitrogen Nutrition of Host Plants

Of the 560 leaf samples belonging to 259 species of green plantsexamined more than 50 per cent of the Angiosperms and 25 percent of the Pteridophytes and Gymnosperms revealed the presenceof N₂-fixing micro-organisms in their phyllosphere. Plants particularlyremarkable in this respect are orchids and several other epiphytes,Scindapsus officinalis, Ficus and cucurbits. Most of the isolatesappear to be biotypes of Klebsiella pneumoniae. The more activestrains fixed more than 5 mg N g^–1 glucose utilized andreduced more than 100 nmol C₂H₂ mg^–1 cell d. w h^–1. The efficacy of the phyllosphere N₂-fixing isolates for N-nutritionof host plants was studied by spraying suspensions of the culturesgrown on N-free media on rice and wheat seedlings. In IR-26rice or Sonalika and Janak wheat grown on soil in wooden flatsor earthenware pots, 22 per cent of the 161 cultures studiedcaused increased height and about three-quarters of the culturesenhanced dry weight by more than 50 per cent; chlorophyll andN-contents were enhanced more than 50 per cent by about halfand two-thirds of the cultures respectively. In N-free sandculture 26 of the 50 promising strains doubled N-content, and30 doubled dry weight of the tested plants. In some cases dryweight, number of grains per panicle, and 1000 grain weightwere increased by 300, 70–83 and 126–158 per centrespectively; N-content of straw and seed was increased three-or fourfold. In several cases the beneficial effects were foundto match closely the performance of plants receiving ammoniumsulphate. Nitrogen-fixing micro-organisms, nitrogen nutrition, phyllosphere, rice, tropical plants, wheat     

Influence of Elevated CO2 and Temperature on the Photosynthesis and Respiration of White Clover Dependent on N2 Fixation

Single clonal plants of white clover (Trifolium repens L) grownfrom explants in a Perlite rooting medium, and dependent fornitrogen on N₂ fixation in root nodules, were grown for severalweeks in controlled environments which provided two regimesof CO₂, and temperature 23/18 °C day/night temperaturesat 680 µmol mol^–1 CO₂, (C680), and 20/15 °Cday/night temperatures at 340 µmol mol^–1 CO₂ (C340)After 3–4 weeks of growth, when the plants were acclimatedto the environmental regimes, leaf and whole-plant photosynthesisand respiration were measured using conventional infra-red gasanalysis techniques Elevated CO₂ and temperature increased ratesof photosynthesis of young, fully expanded leaves at the growthirradiance by 17–29%, despite decreased stomatal conductancesand transpiration rates Water use efficiency (mol CO₂ mol H₂O^–1)was also significantly increased Plants acclimated to elevatedCO₂, and temperature exhibited rates of leaf photosynthesisvery similar to those of C340 leaves ‘instantaneously’exposed to the C680 regime However, leaves developed in theC680 regime photosynthesised less rapidly than C340 leaves whenboth were exposed to a normal CO₂, and temperature environmentIn measurements where irradiance was varied, the enhancementof photosynthesis in elevated CO₂ at 23 °C increased graduallyfrom approx 10 % at 100 µmol m^–1 s^–1 to >27 % at 1170 µmol m^–2 s^–1 In parallel, wateruse efficiency increased by 20–40 % at 315 µmolm^–2 s^–1 In parallel, water use efficiency increasedby 20–40 % at 315 µmol m^–2 s^–1 In parallel,water use efficiency increased by 20–40 % at 315 µmolm^–2 s^–1 In parallel, water use efficiency increasedby 20–40 % at 315 µmol m^–2 s^–1 to approx100 % at the highest irradiance Elevated CO₂, and temperatureincreased whole-plant photosynthesis by > 40 %, when expressedin terms of shoot surface area or shoot weight No effects ofelevated CO₂ and temperature on rate of tissue respiration,either during growth or measurement, were established for singleleaves or for whole plants Dependence on N₂, fixation in rootnodules appeared to have no detrimental effect on photosyntheticperformance in elevated CO₂, and temperature Trifolium repens, white clover, photosynthesis, respiration, elevated CO₂, elevated temperature, water use efficiency, N₂ fixation     

Xylem fluxes of fixed N through nodules of the legume Acacia littorea and haustoria of an associated N-dependent root hemiparasite Olax phyllanthi

Nodulated 1-1.5-year-old plants of Acacia littorea grown inminus nitrogen culture were each partnered with a single seedlingof the root hemiparasite Olax phyllanthi. Partitioning of fixedN between plant organs of the host and parasite was studiedfor the period 4–8 months after introducing the parasite.N fluxes through nodules of Acacia and xylem-tapping haustoriaof Olax were compared using measured xylem flows of fixed Nand anatomical information for the two organs. N₂ fixation duringthe study interval (635 µg N g FW nodules^–1 d^–1)corresponded to a xylem loading flux of 0.20 µg N mm^–2d^–1 across the secretory membranes of the pencycle parenchymaof the nodule vascular strands. A much higher flux of N (4891µg mm^–2 d^–1) exited through xylem at the junctionof nodule and root. The corresponding flux of N from host xylemacross absorptive membranes of the endophyte parenchyma of Olaxhaustorium was 1.15 µg N mm^–1 d^–1, six timesthe loading flux in nodules. The exit flux from haustorium toparasite rootlet was 20.0 pg N mm^–1 d^–1, 200-foldless than that passing through xylem elements of the nodule.Fluxes of individual amino compounds in xylem of nodule andhaustorium were assessed on a molar and N basis. N flux valuesare related to data for transpiration and partitioning of Cand N of the association recorded in a companion paper. Key words: Olax phyllanthi, host-parasite relationships, N flux, Acacia, N₂ fixation     

Are Mistletoes Shade Plants? CO2Assimilation and Chlorophyll Fluorescence of Temperate Mistletoes and their Hosts

Mistletoes usually have slower rates of photosynthesis thantheir hosts. This study examines CO₂assimilation, chlorophyllfluorescence and the chlorophyll content of temperate host–parasitepairs (nine hosts parasitized by Ileostylus micranthus and Carpodetusserratus parasitized by Tupeia antarctica). The hosts of I.micranthus had higher mean annual CO₂assimilation (3.59 ±0.41 µmol m^-2 s^-1) than I. micranthus(2.42 ± 0.20µmol m^-2 s^-1), and C. serratus(2.41 ± 0.43 µmolm^-2 s^-1) showed higher CO₂assimilation than T. antarctica(0.67± 0.64 µmol m^-2 s^-1). Hosts saturated at significantlyhigher electron transport rates (ETR) and light levels thanmistletoes. The positive relationship between CO₂assimilationand electron transport suggests that the lower CO₂assimilationrates in mistletoes are a consequence of lower electron transportrates. When photosynthetic rates, ETR and chlorophyll a /b ratioswere adjusted for photosynthetically active radiation, hostsdid not have significantly higher CO₂assimilation (3.21 ±0.37 µmol m^-2 s^-1) than mistletoes (2.54 ± 0.41µmol m^-2 s^-1), but still had significantly higher ETRand chlorophyll a / b ratios. The electron transport rates,saturating light and chlorophyll a / b ratios of sun leavesfrom mistletoes were similar to host shade leaves. These responsesindicate that in comparison with their hosts, mistletoe leaveshave the photosynthetic characteristics of the leaves of shadeplants. Copyright 2000 Annals of Botany Company CO₂assimilation, photosynthetic active radiation (PAR), chlorophyll fluorescence, electron transport rate (ETR), photochemical quenching (q_p), non-photochemical quenching (q_n), sun and shade leaves, chlorophyll content, Ileostylus micranthus, Tupeia antarctica, New Zealand     

Antitranspirant-induced Heat Stress in the Parasitic Plant Striga hermonthica--A Novel Method of Control

Leaf temperatures (T₁) of the parasitic plant Striga hermonthicaare substantially below those of the air (T_a), [T_a–T₁]reaching 7 ?C at T_a = 40 ?C. This results from high rates oftranspiration and the consequent evaporative cooling of theleaf. Application of an antitranspirant, which mechanicallyimpedes foliar loss of water vapour, reduced transpiration andstomatal conductance by 40% and 57%, respectively, and reduced[T_a– T₁] to 2 ?C at T_a = 40 ?C. The temperature sensitivityof photosynthesis in the host-parasite association differed,the optima (T_opt) being 37.2 and 40.1 ?C for S. hermonthicaand sorghum, respectively. Once Topt had been exceeded in S.hermonthica net photosynthesis declined rapidly, reaching thelethal limit (Tmax) at 42.6 ?C. S. hermonthica is particularlysensitive to high temperatures and antitranspirant-induced overheatingleads to blackening and shrivelling of the leaf after as littleas 4 h at T_a = 40 ?C. Application of an antitranspirant underfield conditions in the Sudan at T_a = 40 ?C resulted in 28%and 67% reductions in transpiration and stomatal conductance,together with a 5 ?C increase in T₁, and subsequent leaf death.In addition to these short-term physiological responses, antitranspirantspraying of the arasite increased the grain and straw yieldof the crop by factors of 3.4 and 2.6, respectively. Antitranspirantsmay have potential use as a method of controlling Striga inthe field. Key words: Striga hermonthica, sorghum, photosynthesis, transpiration, high temperature stress, anti-transpirant     

Apparatus for the Simultaneous Measurement of Water Vapour and Carbon Dioxide Exchanges of Single Leaves

Equipment is described which delivers air with concentrationsof CO₂ and water vapour closely controlled in the ranges 0 to2500 ppm and 5 to 15 mb respectively, at flow rates of up to10 1 min^-1, to each four leaf chambers. The leaf temperatureis controlled to ±0.5 °C and, with a light intensityof 0.3 cal cm^-2 min^-1 visible radiation (0.4 to 0.7 µm)leaf temperature can be maintained at 17.5 °C.The apparatusused to measure the concentration differences between the watervapour and CO₂ entering and leaving the leaf chamber (used tocalculate transpiration, photosynthetic, and respiration rates)is described in detail.Results of tests, which show the necessityfor mounting a fan within the leaf chamber, are reported.Typicallight- and CO₂-response curves are given for kale leaves (Brassicaoleracca var. acephala) and an attempt is made to quantify theerrors in the measurement of photosynthesis and transpiration.     

Can the 15N Dilution Technique be used to Study N2 Fixation in Tropical Tree Symbioses as Affected by Water Deficit?

Three methods were used to study N₂ fixation and effects ofwater deficit on N₂ fixation: C₂H₂ reduction assay (ARA), ¹⁵Ndilution technique and accumulated N content. In addition, ¹⁵Ndilution was calculated both in a traditional way and in a modifiedway, which takes into consideration N and ¹⁵N content for theplants before the experiment started. The three methods wereapplied on the following Rhizobium-symbioses: Acacia albidaDel (Faidherbia albida (Del) A. Chev.) and Leucaena leucocephala(Lam) de Wit., and the Frankia-symbiosis Casuarina equisetifoliaL. The plants wereabout 4-months-old when they were harvested. Nitrogen derived from N₂ fixation in control plants of Acaciaalbida was 54·2 mg as measured with ARA, while it was28·5 mg as measured with the ¹⁵N dilution technique,compared to 30·7 mg calculated as accumulated N. In comparison,L. leucocephala fixed 41·6 mg N (ARA), 53·5 mgN(15N dilution technique) and 56·3 mg N (accumulatedN). The Frankia-symbiosis had fixed 27·4 mg N as measuredby ARA, 8·1 mg N as measured by ¹⁵N dilution techniqueand 12·3 mg N as accumulated N. There were no differencesbetween the estimates based ontraditional and modified waysof calculating ¹⁵N dilution. The immediate effect of water deficit treatment on N₂ fixationwas continuously measured inall species with ARA, which startedto decrease approximately 10 d after the initiation of the treatment,and declined to less than 5% of the initial level after 21–28d. The decrease in the amount of N derived from N₂ fixation wasstudied in L. leucocephala during the period of treatment. Therewas a 26% decrease in amount of N derived from N₂ fixation asresult of water deficit (as measured with ARA), while the decreasewas 23% when measured withboth the ¹⁵N dilution method and asaccumulated N. The three different methods for measuring N₂ fixation and effectsof water deficit on N₂ fixation are discussed. Key words: Acacia albida, ARA, Casuarina equisetifolia, Leucaena leucocephala, ¹⁵N dilution, N₂N fixation, water deficit     

Responses of N2 Fixation-linked Respiration to Host-plant Energy Status in White Clover Acclimated to a Controlled Environment

Ryle, G. J. A., Powell, C. E. and Gordon, A. J. 1988. Responsesof N₂ fixation-linked respiration to host-plant energy statusin white clover acclimated to a controlled environment.—J.exp. Bot. 39: 879–887. Single plants of white clover, acclimated to a controlled environmentand dependent for nitrogen on N₂ fixation in their root nodules,were darkened, defoliated or exposed to enhanced CO₂ levelsto establish the quantitative relationships between the photosynthesisof the host plant and the N₂ fixation metabolism of root nodules. The nodule respiration associated with N₂ fixation (FLR) declinedrapidly to 10–15% of its normal rate following plant darkeningearly in the photoperiod. Darkening at progressively later intervalsduring the photoperiod demonstrated a positive, apparently linearrelationship between duration of illumination and total FLRduring the photoperiod and the following night period. Completeor partial defoliation reduced FLR according to the leaf arearemoved: again, there was a strong positive correlation betweencurrent rate of photosynthesis, whether of defoliated or undefoliatedplants and the FLR of root nodules. Doubling the current rateof photosynthesis, by enhancing CO₂ levels around the shoots,promoted FLR within 1–2 h when plants were stressed bylack of light. However, enhanced CO₂ levels increased FLR onlyslowly over a period of several hours in plants entrained tothe normal growing conditions. It is concluded that, in these plants acclimated to a uniformand favourable controlled environment, the supply and utilizationof photosynthetic assimilate in N₂ fixation was finely balancedand quantitatively linked during a single diurnal period andthat nodule functioning was not depressed by lack of energysubstrate. Key words: White clover, N₂ fixation, photosynthesis.     

Carbon and nitrogen isotope ratios, nitrogen content and heterotrophy in New Zealand mistletoes

The carbon isotope ratio ('¹³C) of New Zealand mistletoes (-29.51ǂ.10‰) and their hosts (-28.89ǂ.12‰) is generally more negative, and shows less difference between mistletoes and their hosts, than found in previous studies. In 37% of the examined pairs, the '¹³C of mistletoes was less negative than that of their hosts. These reversals were not associated with the relative position (proximal or distal) of the host material with regard to the mistletoe. Differences between host and mistletoe tended to be greater on hosts with less negative '¹³C. Both nitrogen content and isotope ratio ('¹⁵N) of the mistletoe leaves were strongly correlated with those of their hosts. Nitrogen contents of mistletoe leaves were similar to those of their hosts at low nitrogen contents but proportionately less on hosts with a high nitrogen content, whereas '¹⁵N of mistletoes was consistently similar to that of their hosts. The '¹³C of mistletoes was related to both host nitrogen content and '¹⁵N, but '¹³C in host tissue was related to neither, suggesting that the mistletoes derived both nitrogen and carbon from their hosts. The '¹³C of both hosts and mistletoes were significantly related to leaf conductance and carbon dioxide concentration but relationships with transpiration and water use efficiency were not significant. In all cases there was no clear separation between the responses of hosts and mistletoes. This may be related to the similarity of stomatal conductance, transpiration and photosynthesis in the studied mistletoes and their hosts and is consistent with the small differences in '¹³C between mistletoes and hosts found in this study. Consequently, the estimation of mistletoe heterotrophy from carbon discrimination is confounded, as the small difference between host and mistletoe carbon discrimination could equally well result from either similarities in photosynthesis and water relations or heterotrophic assimilation of host-derived carbon. The differences between our study and previous studies (which are mostly from seasonally dry or semi-arid to arid environments) may be related to the temperate environment in which these mistletoes grow. Water is freely available so that the mistletoe is able to obtain sufficient water and dissolved nutrients without having to maintain the high transpiration rate and low water potentials that are needed to extract water from a water-stressed host. Similarly, mistletoe photosynthesis is less inhibited by water stress. The physiological similarities between mistletoe and hosts from a temperate environment are reflected in their similar '¹³C values.     

The Relation Between the Nitrogen Concentration and Photosynthetic Capacity of Potato (Solanum tuberosum L.) Leaves

Measurements of the rate of light-saturated photosynthesis (P_max)were made on terminal leaflets of potato plants growing in cropssupplied with 0, 3, 6, 12, 24 and 36 g N m^–2. Measurementswere made between 100 and 154 d after planting. Two types ofleaf were selected—the fourth leaf on the second-levelbranch (L₄, _B1) and the youngest terminal leaflet that was measurable(L_YM). Later, the total nitrogen concentration of each leaflet(N_L) was measured. A linear regression between P_max and N_L,common to both leaf positions, explained 68.5% of the totalvariation. With L₄, _B1 leaves there was a significant improvementin the proportion of variation explained when regressions withseparate intercepts and a common slope were fitted to individualfertilizer treatments. These results suggest that an increasingproportion of leaf nitrogen was not associated with the performanceof the photosynthetic system with increasing nitrogen supply.This separation between nitrogen treatments was not as clearfor L_YM leaves. Stomatal conductance to transfer of water vapourwas neither influenced by leaf position nor directly by nitrogensupply. Rather conductance declined in parallel with the declinein photosynthetic capacity. Solanum tuberosum, potato, nitrogen, photosynthesis, stomatal conductance, leaf     

Haustorial Development and Growth Benefit to Seedlings of the Root Hemiparasitic Tree Nuytsia floribunda(Labill.) R.Br. in Association with Various Hosts

Dry matter gains and haustorial production of pot-cultured seedlingsof Nuytsia floribunda were assessed after a 12 month periodof association singly with each of a range of potential woodyhost species. One species,Adenanthos cygnorum , of similar sizeto most parasitized hosts, served as measure of response ofNuytsia in a non-benefiting situation. Rated on this basis,all 23 parasitized hosts elicited greater mean dry weights ofNuytsia than when on Adenanthos, and seven of these instanceswere highly significant. Numbers and weights of penetratingand presumably functional haustoria formed on a host were broadlycorrelated with growth benefit to Nuytsia, but there were notableinstances of unusually poor or great benefit from a host relativeto the complement of haustoria involved. Experiments in whichhaustoria-bearing associations of Nuytsia partnered with nodulatedAcacia hosts (Acacia acuminata and A. cyclops) were fed¹⁵N₂showedsignificant transfer of¹⁵N to the parasite, but failed to determinewhether the label had been acquired through haustoria or directlyby Nuytsia roots following turnover of nodule and root residuesof the host in the rooting medium. A parallel study using theunusual non-protein amino acid, djenkolic acid, as a markerof benefit from the djenkolic acid-containing host A. cyclops,showed appearance and progressive build-up of the compound infoliage of Nuytsia over a 6 month period after partnering thespecies in pot culture. Presence of the compound at final harvestin xylem sap of both partners but not in soil solution of thecultures strongly indicated xylem transfer via haustoria asthe principal avenue for N benefit to the parasite. Resultsare discussed in relation to a recent evaluation of haustorialstructure and functioning of N. floribunda. Copyright 2000 Annalsof Botany Company Root hemiparasite, Nuytsia, Loranthaceae, growth benefit, haustorial production, nitrogen transfer from hosts     

The Influence of Temperature on Photosynthesis and Transpiration in ten Temperate Grass Varieties Grown in four Different Environments

The influence of temperature on photosynthesis and transpirationwas studied in ten varieties of Lolium perenne, L. multiflorum,Dactylis glomerata, and Festuca arundinacea from three climaticorigins grown in three different controlled environments (15?C, 72 W m^-2 visible irradiation, 16-h photoperiod; 25 ?C, 72W m^-2 visible irradiation, 16-h photoperiod; and 25 ?C, 180W m^-2 visible irradiation, 16-h photoperiod) and in the glasshousein July/August. The optimum temperature for photosynthesis was influenced primarilyby growth environment; growth at low temperature (15 ?C) resultedin a low optimum temperature, which differed little from varietyto variety. The maximum CO₂-exchange rate was influenced bygrowth environment and by variety. Within a variety, plantsgrown at higher light intensity or lower temperature had a greaterCO₂-exchange rate. Seven varieties showed a negative correlationbetween the optimum leaf temperature and the maximum CO₂-exchangerate. Activation energies for photosynthesis were influencedby growth environment only. There were marked varietal differences in the values of leafresistances (r_a + r_t) obtained from transpiration data at theoptimum leaf temperature for CO₂ exchange. In Lolium, and Dactylisthe Mediterranean varieties had higher leaf resistances thanthe Northern varieties with the maritime varieties intermediate.In general the Dactylis varieties had higher resistances thanthe corresponding Lolium and Festuca varieties. Only at highgrowth temperatures was (r_a+r_l) insensitive to temperature;otherwise an activation energy of about 10 kcal/mole was observed.A negative correlation was found between mean varietal diffusionresistances (r_a+r_l), and corresponding maximum CO₂-exchangerates.