Germination stimulants produced by Vigna unguiculata Walp cv Saunders Upright

A germination stimulant, alectrol, for the seeds of the angiospermous root parasites Alectra vogelii Benth. and Striga gesnerioides (Willd.) Vatke has been isolated from root exudates of its genuine host plant Vigna unguiculata Walp cv Saunders Upright. Its spectroscopic data lead to a chemical structure closely related to (+)-strigol. The compound is more active than strigol in stimulating seeds of the respective parasites, Alectra vogelii and Striga gesnerioides.Abbreviations AV Alectra vogelii - OA Orobanche aegyptiaca - SA Striga asiatica - SG Striga gesnerioides - SH Striga hermonthica - TFA trifluoroacetic acid     

The Maize Glutamine Synthetase GS1- Gene is Preferentially Expressed in Kernel Pedicels and is Developmentally-Regulated

The pedicel region of Zea mays kernels contains a unique formof maize glutamine synthetase (GS), GS_pl. RNA blot analysisusing GS gene-specific probes revealed that the expression ofthe GS.₂ gene was specific to the pedicel and that it increasedin the kernels during development. This pattern of the maizeGS.₂ gene expression is consistent with the tissue specificityof the GS_pl protein and suggests that it encodes the GS_pl isoformof maize GS. (Received September 16, 1997; Accepted January 19, 1998)     

Striga seed-germination activity of root exudates and compounds present in stems of Striga host and nonhost (trap crop) plants is reduced due to root colonization by arbuscular mycorrhizal fungi

Root colonization by arbuscular mycorrhizal (AM) fungi reduces stimulation of seed germination of the plant parasite Striga (Orobanchaceae). This reduction can affect not only host plants for Striga, resulting in a lower parasite incidence, but also false hosts or trap crops, which induce suicidal Striga seed germination, thereby diminishing their effectiveness. In order to better understand these AM-induced effects, we tested the influence of root colonization by different AM fungi on the seed-germination activity of root exudates of the Striga hermonthica nonhost plants cowpea and cotton on S. hermonthica. We also tested the effect of AM fungi on the seed-germination activity of the Striga gesnerioides host plant cowpea on S. gesnerioides. Moreover, we studied whether mycorrhization affects the transport of seed-germination activity to above-ground plant parts. Mycorrhization not only resulted in a lower seed germination of S. gesnerioides in the presence of root exudates of the S. gesnerioides host cowpea but also seed germination of S. hermonthica was also lower in the presence of root exudates of the S. hermonthica nonhosts cowpea and cotton. Downregulation of the Striga seed-germination activity occurs not only in root exudates upon root colonization by different AM fungi but also in the compounds produced by stems. The lowered seed-germination activity does not appear to depend on the presence of seed germination inhibitors in the root exudates of mycorrhizal plants. The implication for Striga control in the field is discussed.     

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     

Structural requirements of strigolactones for germination induction and inhibition of Striga gesnerioides seeds

Key message

Structure–activity relationship studies of strigolactones and Striga gesnerioides seed germination revealed strict structural requirements for germination induction and a new function of the plant hormones as germination inhibitors.

Abstract

Stereoisomers of the naturally occurring strigolactones, strigol, sorgolactone, orobanchol, sorgomol and 5-deoxystrigol, 36 in total, were prepared and screened for the ability to induce and/or inhibit the germination of Striga hermonthica and Striga gesnerioides seeds collected from mature plants that parasitized on sorghum and cowpea, respectively. All of the compounds induced S. hermonthica seed germination, albeit displayed differential activities. On the other hand, only a limited number of the compounds induced significant germination in S. gesnerioides, thus indicating strict structural requirements. Strigolactones inducing high germination in S. gesnerioides induced low germination in S. hermonthica. Strigolactones with the same configuration at C3a, C8b and C2′ as that in 5-deoxystrigol (9a) induced high germination of S. hermonthica seeds, but most of them inhibited the germination of S. gesnerioides. The differential response of S. gesnerioides to strigolactones may play an important role in the survival of the species. However, the compounds could be used as means of control if mixed cropping of cowpea and sorghum is adopted.     

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     

Striga gesnerioidesParasitising Cowpea: Development of Infection Structures and Mechanisms of Penetration

Striga gesnerioides(Scrophulariaceae) is an obligate hemiparasiticangiosperm, that infects the roots of cowpea (Vigna unguiculata).The development ofStrigaseedlings was characterized after transferto the surface of cowpea roots using cryo-scanning electronmicroscopy. Contact with the surface of the root caused distinctmorphological changes in theStrigaradicles. Radicle elongationceased 8 h after contact with host roots and radicle hairs developed.These hairs adhered to the surface of the host root. Early stagesof root penetration, i.e. prior to connection to the host vascularsystem were examined by light microscopy and transmission electronmicroscopy to determine the mechanisms by whichStrigainvadesroot tissue. Cowpea roots were penetrated byS. gesnerioidesradiclesafter 48 h. Host cells immediately surrounding the invadingStrigawerecompressed, but other host cells were not affected. Immunocytochemistry,using monoclonal antibodies JIM 5 and JIM 7, showed thatStrigaadvancedintercellularly through the root tissues without extensive removalof pectins at the site of penetration.Copyright 1998 Annalsof Botany Company Striga gesnerioides,Vigna unguiculata, mechanisms of penetration, host-parasite interactions, electron microscopy, immunocytochemistry.     

Striga hermonthica decreases photosynthesis in Zea mays through effects on leaf cell structure

Maize seedlings were grown in pots either with or without preconditionedseeds of the parasitic weed, Striga hermonthica. After between4 and 8 weeks, net photosynthesis in the leaves of maize plantsinfected with Striga decreased compared to leaves of uninfectedcontrol plants. The activities of four enzymes of photosyntheticmetabolism were, however, little affected by infection. A pulse-chaseexperiment using ¹⁴CO₂ showed that C₄ acids were the main earlyproducts of assimilation even when the rate of photosynthesiswas much decreased by infection, but more radio-activity appearedin glycine and serine than in leaves of healthy maize plants.Leaves of infected maize required longer to reach a steady rateof photosynthesis upon enclosure in a leaf chamber than leavesof uninfected plants after similar treatment. Electron microscopy of transverse sections of the leaves ofinfected maize indicated that the cell walls in the bundle sheathand vascular tissue were less robust than in leaves of healthyplants. The results suggest that infection with Striga causesan increase in the permeability of cell walls in the bundlesheath, leakage of CO₂ from the bundle sheath cells and decreasedeffectiveness of C₄ photosynthesis in host leaves. Key words: Zea mays, Striga hermonthica, photosynthesis, photorespiration, enzyme activity     

The Effects of Compatible Solutes on the Heat Stability of Glutamine Synthetase from Chickpeas Grown under Different Nitrogen and Temperature Regimes

The activity of glutamine synthetase (GS) extracted from chickpeasgrown under different nitrogen and temperature regimes was examined.Cytosolic GS (GS₁) was much less temperature sensitive thanchloroplastic GS (GS₂). There was no change in the ratio ofthe two isoforms from plants with different growth temperatures.Specific activity of GS appeared to be independent of growthtemperature and was little affected by nitrogen supply. In vitro studies were performed on the heat stability of GS₂in the presence of glycine betaine, proline, and myoinositol.The latter was most effective in preventing loss of enzyme activityat high temperatures. Glycine betaine was also effective asa heat protectant but enzyme activity in the presence of prolinewas lower than the control at temperatures above 30 ?C. Theimportance of myo-inositol protection of metabolism is discussedin relation to the accumulation in chickpeas of another cyclitol,pinitol. Key words: Chickpeas, high temperature stress, glutamine synthetase     

Physiology of the parasitic association between maize and witchweed (Striga hermonthica): is ABA involved?

Striga hermonthica (Del.) Benth. is an obligate hemiparasiticangiosperm which can cause severe losses of yield in cerealcrops in the semi-arid tropics. The effects of this parasiteon the growth and stomatal conductance of three varieties ofmaize (Zea mays L.) during the first 6 weeks of the associationhave been studied. From 24 d after planting (DAP), infectedplants were significantly shorter than uninfected controls.When the plants were harvested 45 DAP, infected plants had fewerfully expanded leaves, less leaf biomass and less pseudo-stembiomass than uninfected controls. However, the parasitized plantshad more root biomass and hence a higher root:shoot ratio thanuninfected controls. The stomatal conductance of infected hostswas severely inhibited by comparison with that in uninfectedplants. The possibility that abscisic acid (ABA) may be involved inthe regulation of the parasitic association was investigated.ABA concentrations in leaf tissue of maize (cv. Cargimontana)and S. hermonthica were determined by radioimmunoassay. Whilethere was a difference between cultivars in the extent of theresponse, the concentrations of ABA were significantly higherin infected maize plants than in the uninfected controls. InS. hermonthica, leaf tissue ABA concentration was found to bean order of magnitude higher than in the host leaf tissue. Detachedleaves of S. hermonthica which were dehydrated at room temperatureuntil they had lost 10–20% of their fresh weight containedthree times the ABA concentration of control leaves. This suggeststhat leaves of S. hermonthica can synthesize or re-mobilizeABA in response to water deficit. It is not yet known whetherthis contributes to the higher concentration in infected hosts,but the results suggest that ABA has a role in this parasiticassociation. Key words: Striga hermonthica, abscisic acid, growth, parasitic angiosperm, stomatal conductance     

Development of Photorespiration during Chloroplast Biogenesis in Wheat Leaves

Tobin, A. K., Sumar, N., Patel, M., Moore, A. L. and Stewart,G. R. 1988. Development of photorespiration during chloroplastbiogenesis in wheat leaves.—J. exp. Bot. 39: 833–843. The rate of light-dependent ammonia accumulation in L-methioninesulphoximine (MSO: glutamine synthetase inhibitor)-treated wheat(Triticum aestivum L. cv. Maris Huntsman) primary leaf sectionsincreased with mesophyll cell maturity. Ammonia production inthe more mature sections (beyond 2.0 cm from the basal meristem)was inhibited by elevated CO₂ concentrations and by incubationwith 10 mol m^–3 pyrid-2-yl hydroxymethane sulphonate (HPMS).In contrast, the low levels of ammonia which accumulated inthe immature sections (0 to 2.0 cm from the base) were unaffectedby such treatments. This indicates that the ammonia producedin mature wheat leaf sections is of photorespiratory originand that the capacity of this pathway increases with mesophyllcell and chloroplast development. Rates of CO₂-dependent oxygenevolution by leaf sections (under saturating CO₂) increasedin parallel with ammonia production. Levels of endogenous nitratewere relatively high and increased from 5.15 mol x 10^–13mesophyll cell^–1 in meristematic cells to 6.6 mol x 10^–12mesophyll cell^–1 in mature tissue. There was no significantchange in leaf nitrate level during 30 min light incubationof the wheat leaf sections, indicating that the majority ofthe nitrate was metabolically inactive and stored in the vacuole.Activities of key enzymes of photorespiration (glutamine synthetase,glycollate oxidase), nitrogen metabolism (nitrate reductase,glutamate dehydrogenase, glutamine synthetase) and mitochondrialrespiration (cytochrome oxidase), showed specific and distinctpatterns of development during leaf growth. Chloroplast glutaminesynthetase (GS₂) and peroxisomal glycollate oxidase developedin apparent synchrony with the major increase in activity occurringin regions beyond4.0 cm from the leaf base, i.e. where photorespirationwas developing. Cytosolic glutamine synthetase (GS₁) and nitratereductase (in vivo) activities were identical throughout leafgrowth, reaching maximum rates at 4.0 cm from the base and thenremaining constant. Activities of the mitochondrial enzymesglutamate dehydrogenase (GDH) and cytochrome oxidase were highin meristematic cells and increased in parallel, attaining amaximum towards the leaf tip. This indicated a respiratory,as opposed to a photorespiratory, role for GDH in wheat leafmetabolism. The evidence for controlled, co-ordinated synthesisof pathway enzymes at specific stages of organelle biogenesisis discussed. Key words: Photorespiration, organelle biogenesis     

STUDIES ON THE GERMINATION OF SEEDS OF THE ROOT PARASITES,ALECTRA VOGELII AND STRIGA GESNERIOIDES. I. ANATOMICAL CHANGES IN THE EMBRYOS

Anatomical changes in the radicle and shoot meristems of embryos of germinating seeds of the obligate root parasites, Alectra vogelii and Striga gesnerioides were studied. When germination of seeds was stimulated by cowpea (Vigna unguiculata) root exudate, growth occurred mainly in the radicular pole of embryos and minimally in the plumular pole, resulting in seedlings with elongated radicles. Maximum radicle elongations of about 3 mm in A. vogelii and 2 mm in S. gesnerioides were recorded during a period of 8 and 11 days, respectively. Analysis of the radicular tip during the course of seed germination revealed that the activity of the meristematic tissue progressively decreased until it completely disappeared. When germinated seeds were cultured on nutrient agar media, the radicle meristem of A. vogelii continued to grow producing a normal root with a root cap. On the other hand, the radicles of cultured S. gesnerioides seeds elongated only slightly before meristematic activity ceased. During continued growth of seedlings of both species on agar media, lateral roots whose tips had typical angiosperm root topography, were initiated from the radicle.     

Evidence for a cytosolic-dependent light induction of chloroplastic glutamine synthetase during greening of etiolated rice leaves

During the greening of etiolated rice leaves, total glutamine synthetase activity increases about twofold, and after 48 h the level of activity usually observed in green leaves is obtained. A density-labeling experiment with deuterium demonstrates that the increase in enzyme activity is due to a synthesis of the enzyme. The enhanced activity obtained upon greening is the result of two different phenomena: there is a fivefold increase of chloroplastic glutamine synthetase content accompanied by a concommitant decrease (twofold) of the cytosolic glutamine synthetase. The increase of chloroplastic glutamine synthetase (GS₂) is only inhibited by cycloheximide and not by lincomycin. This result indicates a cytosolic synthesis of GS₂. The synthesis of GS₂ was confirmed by a quantification of the protein by an immunochemical method. It was demonstrated that GS₂ protein content in green leaves is fivefold higher than in etiolated leaves.Abbreviations AbH heavy chain of antibodies - AbL light chain of antibodies - AP acid phosphatase - CH cycloheximide - G6PDH glucose-6-phosphate dehydrogenase - GS glutamine synthetase - GS₁ cytosolic glutamine synthetase - GS₂ chloroplastic glutamine synthetase - LC lincomycin - NAD-MDH NAD malate dehydrogenase - NADP-G3PDH NADP glyceraldehyde-3-phosphate dehydrogenase     

Root nitrogen concentration of sorghum above 2% produces least Striga hermonthica seed stimulation

A series of pot and laboratory experiments was carried out to assess the effects of N status of sorghum roots and timing of N application (as NH₄NO₃) on the germination of Striga hermonthica seeds. Root N concentrations varied from 10 to 26 mg N g^?1. The cut root and the root exudates technique used in assaying S. hermonthica seed germination gave similar results. However, the cut root technique was easier to handle and more discriminating at low germination levels. S. hermonthica seed germination per unit sorghum root mass followed a broken‐stick model. It decreased with increasing root N concentrations, reaching lowest levels at a root N concentration of 19.5 mg N g^?1, after which no further reduction occurred. It was not possible to reduce S. hermonthica seed germination to a zero level. Timing of N application influenced the time a higher N concentration is reached, not the S. hermonthica seed germination. Both timing and rate of N application are important in maintaining root N concentrations above 19.5 mg N g^?1, thereby potentially reducing S. hermonthica germination in the field. Translation of results to reductions in infection levels and yield losses is hampered by density‐dependent relations after the S. hermonthica germination stage.     

Carbon isotope ratios demonstrate carbon flux from c(4) host to c(3) parasite

Carbon isotope ratios of mature leaves from the C₃ angiosperm root hemiparasites Striga hermonthica (Del.) Benth (−26.7‰) and S. asiatica (L.) Kuntze (−25.6‰) were more negative than their C₄ host, sorghum (Sorghum bicolor [L.] Moench cv CSH1), (−13.5‰). However, in young photosynthetically incompetent plants of S. hermonthica this difference was reduced to less than 1‰. Differences between the carbon isotope ratios of two C₃-C₃ associations, S. gesnerioides (Willd.) Vatke—Vigna unguiculata (L.) Walp. and Oryza sativa L.—Rhamphicarpa fistulosa (Hochst.) Benth differed by less than 1‰. Theoretical carbon isotope ratios for mature leaves of S. hermonthica and S. asiatica, calculated from foliar gas exchange measurements, were −31.8 and −32.0‰, respectively. This difference between the measured and theoretical δ¹³C-values of 5 to 6‰ suggests that even in mature, photosynthetically active plants, there is substantial input of carbon from the C₄ host. We estimate this to be approximately 28% of the total carbon in S. hermonthica and 35% in S. asiatica. This level of carbon transfer contributes to the host's growth reductions observed in Striga-infected sorghum.     

Variation of Seed-coat Ornamentation in Striga hermonthica (Scrophulariaceae)

The surface features of the seed of the parasitic floweringplant Striga hermonthica were examined with the scanning electronmicroscope. The details of ornamentation were constant on seedsfrom one plant but varied within and between populations andare probably due to out-breeding. The variation was not relatedto geographical origin or to host-preference. Testa patternssimilar to those reported for other Striga species were foundin the samples. Striga hermonthica, Scrophulariaceae: Rhinanthoideae, hemi-parasite, testa, seed-coat, scanning electron microscope, genetic variation     

HowStriga Parasitizes its Host: a TEM and SEM Study

The cellular contact betweenStriga hermonthica andStriga asiaticaand their hosts,Zea mays andSorghum bicolor , was investigatedby light, transmission electron and scanning electron microscopy.The xylem connections between parasites and hosts involve veryspecific, clustered intrusions into the host's water conductingelements, predominantly into the large vessel elements. A singlehaustorial cell can penetrate a host vessel element with morethan one intrusion. All intrusions become covered by an additionalelectron-opaque wall layer. During subsequent differentiation,a dissolution of specific wall parts of the cell intrusionsoccurs so that open, cup- or trunk-like structures result. Thevessel-like host contact can comprise up to five openings withina single intrusion. Concomitantly, the intrusions and the haustorialcells to which they belong lose their protoplasts and transforminto elements which take up water. The walls of the haustorialcells and both wall parts of their appendages become stronglylignified. The water and nutrient absorbing structures insertedinto the host vessel are named ‘oscula’. Withinthe whole haustorial complex of bothStriga species no phloemelements were detected. Translocation of substances from hostto parasite are briefly discussed. Striga hermonthica ; Striga asiatica ; haustorial anatomy; xylem contact; osculum     

Xylem exudation is related to nitrate assimilation pathway in detopped maize seedlings: use of nitrate reductase and glutamine synthetase inhibitors as tools

The xylem exudation of detopped 7-d-old seedlings of Zea maysL. doubled when KCI was present in the root medium comparedto seedlings maintained on water. It was further enhanced whenKCI was replaced by nitrogen compounds such as nitrate, ammoniumand glutamine. The role of the nitrate assimilation pathwayon the enhancement of xylem exudation rate was investigatedusing tungstate, an inhibitor of nitrate reductase (NR) activity,and phosphinothricin or methionine sulphoximine, inhibitorsof glutamine synthetase (GS) activity. The sap levels of NO₃^–,NH₄⁺, glutamine, and asparagine was used to ascertain the invivo inhibition of both enzymes. The tungstate effects werealso checked by measuring leaf in vitro NA activity and NR proteincontent. Xylem exudation rate of detopped seedlings fed withKNO₃ decreased when the nitrate assimilation pathway was blockedeither at the NR or at GS sites. This decrease was preventedwhen urea (acting as NH₄⁺ supply) was given simultaneously withtungstate. KNO₃ does not act directly on exudation, but throughthe involvement of NH₄⁺. The involvement of glutamine was alsoshown since GS inhibition resulted in a cancellation of theenhancing effect of KNO₃ on exudation. As change of exudationrate was not linked to change in sap osmolarity, it is assumedthat the assimilation chain could modify root water conductance.The role of glutamine was discussed. Key words: Exudation, maize, nitrate, conductance, NR, GS     

Appearance of a novel form of plant glutamine synthetase during nodule development in Phaseolus vulgaris L.

The activities of glutamine synthetase (GS), nitrogenase and leghaemoglobin were measured during nodule development in Phaseolus vulgaris infected with wild-type or two non-fixing (Fix^-) mutants of Rhizobium phaseoli. The large increase in GS activity which was observed during nodulation with the wild-type rhizobial strain occurred concomitantly with the detection and increase in activity of nitrogenase and the amount of leghaemoglobin. Moreover, this increase in GS was found to be due entirely to the appearance of a novel form of the enzyme (GS_n1) in the nodule. The activity of the form (GS_n2) similar to the root enzyme (GS_r) remained constant throughout the experiment. In nodules produced by infection with the two mutant strains of Rhizobium phaseoli (JL15 and JL19) only trace amounts of GS_n1 and leghaemoglobin were detected.Abbreviations DEAE-Sephacel diethylaminoethyl-Sephacel - GS glutamine synthetase     

Growth and carbon allocation in Pennisetum typhoides infected with the parasitic angiosperm Striga hermonthica

Abstract Growth and gas exchange measurements are used in conjunction with a carbon balance model to describe the millet (Pennisetum typhoides)–witchweed (Striga hermonthica) host—parasite association. Striga hermonthica reduces the growth of millet by 28% and radically alters the architecture of infected plants. Whilst grain yield and stem dry weight are reduced (by 80 and 53%, respectively), leaf and root growth are stimulated (by 41 and 86%, respectively). The difference in production between infected and uninfected millet plants can be accounted for by two processes: first, export of carbon to the parasite (accounting for 16% of the dry weight not gained); and second, parasite-induced reductions in host photosynthesis (accounting for 84% of the dry weight not gained). Striga hermonthica is dependent on carbon exported from the host, since the plant has low rates of photosynthesis coupled with high rates of respiration. The carbon balance model suggests that in mature S. hermonthica plants parasitic on millet, 85% of the carbon is host-derived. Carbon fluxes are also estimated using δ¹³C measurements, since S. hermonthica is a C₃ plant parasitizing a C₄ host. In conjunction with gas exchange measurements, these suggest that in root, stem and leaf of S. hermonthica, 87, 70 and 49% of carbon is hostderived, respectively.