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     

Structure and Development of the Primary Haustorium in Alectra vogelii Benth. (Scrophulariaceae)

Anatomical observations were made on the structure and developmentof the primary haustorium of Alectra vogelii. Its developmentinvolves a mutual aggressive growth of both the host and parasitetissues resulting in the formation of a very large and complextuberous organ. One of the host tissues whose growth is stimulatedby parasite infection is the pericycle whose cells divide repeatedlyand grow around and within the parasite haustorial cortex. Fromvarious points of the proliferating host pericycle, roots becomeinitiated and eventually the entire surface of the haustoriumbecomes covered with these roots. We have referred to them as‘haustorial roots’, a term which we have re-examinedand redefined. True xylary connections are established not onlybetween the parasite and the host root but also between theparasite and these ‘haustorial roots’. The uniquedevelopment of primary haustorium and ‘haustorial roots’in A. vogelii is discussed in relation to the development ofprimary haustoria in other root parasites.     

Structure and Development of the Mature Secondary Haustorium in Alectra vogelii Benth

Anatomical investigations were carried out on the structureand development of the mature secondary haustorium in Alectravogelii growing on Arachis hypogaea or Vigna unguiculata. Followingthe formation of the young secondary haustorium, both the cambiumand pericycle of the host root directly opposite the young secondaryhaustorium are stimulated to divide and form new tissues andorgans including haustorial roots. Further proliferations ofthe host root pericycle and the haustorial cortex give riseto a large, tuberous and complex mature secondary haustoriumwithin which the tissues of the host and parasite remain inintimate contact forming a perfect graft union with a wide zoneof contact. Apart from the haustorial axial xylcm strand whichnormally connects the xylem of the parasite secondary root withthat of the host, direct xylary connections are also establishedbetween the axial xylem of the haustorium and the xylem of thehaustorial roots. The entire surface of the mature secondaryhaustorium of Alectrais covered with these haustorial rootsas was previously observed in its mature primary haustorium. Alectra vogelii Benth, secondary haustorium, haustorium, haustorial roots, root parasite, hemiparasitism, Arachis hypogaea, Vigna unguiculata     

Establishment, Structure and Morphology of the Tuber of Balanophora

During germination of the ‘seed’ of Balanophora,endosperm cells at the radicular pole grow out as tubular structuresand anchor the ‘seed’ to the host rootlet. The radiculartier of cells of the embryo elongate as primary haustorial tubesand establish contact with the host root vasculature. A secondaryhaustorium arises from a meristem adjoining the primary haustorium.The remainder of the embryo contributes to the tuber proper. Host parenchyma in the immediate vicinity of the primary haustoriumreverts to meristematic activity. Some of the derivatives matureas perforate tracheary cells. The remainder, retaining meristematicactivity, squeeze themselves between secondary haustorial cellsand together initiate a composite conducting strand, which repeatedlydichotomizes as the tuber grows. The conducting strand of Balanophora is looked upon as the equivalentof combined adventitious root system of parasite and host. Theremaining part of the tuber is equivalent to the shoot. Balanophora, tuber, morphology, host-parasite relations, parasite     

Morphology and Anatomy of the Haustorium of the Root Hemiparasite Olax phyllanthi (Olacaceae), with Special Reference to the Haustorial Interface

Gross morphology and internal structure of haustoria of Olaxphyllanthi are described in parasitism with a range of hosts,including roots of woody and herbaceous dicotyledons and certainmonocotyledons, and occasional instances of autoparasitism andhaustorial formation on monocotyledon rhizomes. Successful penetrationto xylem occurs on virtually all hosts across broad diameters,ages and anatomies of host root, but anatomical impedimentsto haustorial establishment and penetration are recorded forcertain host taxa. Each haustorium is a comparatively simpleand ephemeral structure. Its developing sucker (endophytic regionof the haustorium) spreads laterally around the surface of thehost xylem, yet never completely encircles the host stele. Damageto hosts is minimal and secondary thickening (of hosts) continueson the side of a host root opposite to a haustorium. The haustorialsucker lacks phloem and its interface with host xylem is comprisedalmost entirely (more than 98.7%) of parenchyma. The few terminatingtracheids at an interface lie in very close proximity to oroccasionally directly against exposed xylem vessels, but lumento lumen continuity between tracheary elements of the partnersis not achieved. Three dimensional reconstructions based onserial transverse sectioning indicate that well defined filesof tracheids connect back from an interface to the core of graniferoustracheary elements in the external body of the haustorium, andthence to the xylem of the parent parasite root. The findingsare discussed in relation to existing studies on haustorialanatomy. Root parasite, Olacaceae, haustorial anatomy, host specificity     

Initiation, Development and Structure of the Primary Haustorim in Striga gesnerioides (Scrophulariaceae)

A light-microscopic study is reported on the initiation, establishmentand structure of the primary haustorium of Striga gesnerioideson the host, cowpea (Vigna unguiculata). The radicular apexof the germinated parasite seed dissolves its way through thehost root cortex to the stele. Thus, it is converted into aprimary haustorium. Some of the haustorial front-line cellsin contact with the host endodermis penetrate into the steleand make contact with the xylem vessels. Differentiation ofthese haustorial cells into xylem vessels occurs and extendsbackwards through the median axial region of the haustorialtract in the host cortex to connect with the conductive xylemof the radicle outside the host root. Subsequently the parasite'splumule develops into a leafy shoot. On penetrating the steleof the host, the haustorium stimulates cell division in thehost pericycle whose triggered proliferation together with expansionof the parasite haustorial tissues result in the formation ofa large, tuberous primary haustorium. At various points of thehost-parasite interface, differentiation of xylem elements occurs,presumably maximizing nutrient transfer from host to parasite.In spite of this, many proliferated host cells at the interfaceremain apparently meristematic showing densely-stained cytoplasmand prominent nuclei.     

Does legume nitrogen fixation underpin host quality for the hemiparasitic plant Rhinanthus minor?

The high quality of leguminous hosts for the parasitic plantRhinanthus minor (in terms of growth and fecundity), comparedwith forbs (non-leguminous dicots) has long been assumed tobe a function of the legume's ability to fix atmospheric nitrogen(N) from the air and the potential for direct transfer of compatibleamino compounds to the parasite. Using associations betweenRhinanthus minor and Vicia faba (Fabaceae) that receive N eitherexclusively via symbiotic associations with rhizobia supplyingorganic N fixed from N₂ or exclusively through the supply ofinorganic nitrate to the substrate, the underlying reasons forthe quality of legumes as hosts for this parasite are unravelled.It is shown that sole dependence of the host, V. faba, on Nfixation results in lower growth of the attached parasite thanwhen the host is grown in a substrate supplied exclusively withinorganic N. In contrast, the host plants themselves achieveda similar biomass irrespective of their N source. The physiologicalbasis for this is investigated in terms of N and abscisic acid(ABA) partitioning, haustorial penetration, and xylem sap aminoacid profiles. It is concluded that legume N fixation does notunderpin the quality of legumes as hosts for Rhinanthus butrather the well-developed haustorium formed by the parasite,coupled with the lack of defensive response of the host tissuesto the invading haustorium and the presence of sufficient nitrogenouscompounds in the xylem sap accessible to the parasite haustoria,would appear to be the primary factors influencing host qualityof the legumes. Key words: ABA, haustorium, legume, nitrogen fixation, nodules, parasitic plant Received 14 November 2007; Revised 7 January 2008 Accepted 8 January 2008     

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     

The Initiation of the Secondary Haustorium in Alectra vogelii Benth

Anatomical studies were carried out on initiation of the secondaryhaustorium in Alectra vogelii, a root parasite of leguminouscrops in Nigeria. In both the normal and self-haustorium, theformation of the haustorial initial on the parasite root soonafter initial contact between the host and parasite roots isfollowed by the penetration of the host root by the haustorium.Specialized penetrating cells (intrusive cells) at the haustorialfront prise apart and loosen the host root cortical cells, whichlater become digested. Through the same processes, a few ofthese columnar intrusive cells at the haustorial front piercethe endodermis to make contact with the xylem of the host root.Thereafter, a true conductive bridge consisting of short, isodiametric,reticulate vessel elements is established between the parasiteand host roots through the secondary haustorium. No pholem tissuewas observed in the connection. There is a close similaritybetween the mode of initiation of the secondary haustorium ofAlectra vogelii and that previously described for its primaryhaustorium. Alectra vogelii Benth, haustorium, self-haustorium, root parasite, hemiparasitism, Vigna unguiculata, Arachis hypogaea     

Ultrastructure of the haustorial interface and apoplastic continuum between host and the root hemiparasiteOlax phyllanthi (Labill.) R. Br. (Olacaceae)

Summary Structural features of haustorial interface parenchyma of the root hemiparasiteOlax phyllanthi are described. Walls contacting host xylem are thickened non-uniformly with polysaccharides, not lignin, and show only a thin protective wall layer when abutting pits in walls of host xylem vessels or tracheids. Lateral walls of interface parenchyma exhibit an expanded middle layer of open fibrillar appearance, sometimes with, but mostly lacking adjoining layers of dense wall material. Free ribosomes and rough endoplasmic reticulum are prominent and occasional wall ingrowths present. Experiments involving transpirational feeding of the apoplast tracers lanthanum nitrate or uranyl acetate to host roots cut below haustorial connections, indicate effective apoplastic transfer from host to parasite root via the haustorium. Deposits of the tracers suggest a major pathway for water flow through host xylem pits, across the thin protective wall layer, and thence into the haustorium via the electronopaque regions of the terminal and lateral walls of the contact parenchyma. Graniferous tracheary elements and walls of parenchyma cells of the body of the haustorium appear to participate in tracer flow as do walls of cortical cells, stele parenchyma and xylem conducting elements of the parasite root, suggesting that both vascular and non-vascular routes are involved in extracytoplasmic transfer of xylem sap from host to parasite. The Casparian strip of the endodermis and the suberin lamella of the exodermis of theOlax root act as barriers to flow within the system.     

Anatomical Studies of Haustorium Ontogeny and the Remarkable Mode of Penetration of the Haustorium in Nuytsiafloribunda (Labill.) R. Br.

The development and structure of secondary haustoria of Nuytsia floribunda are described and compared with other Santalalean haustoria. After establishing contact with the host root, cortical folds of the haustorium grow around the root in separate directions and fuse forming a ring around it. At an early stage of development, meristematic tissue differentiates in the interior proximal part of the haustorium. Zones of collapsed layers are present in the outer cortical region. Subsequently, in the proximal part, two vascular cores, two lysigenous cavities and extensive masses of sclerenchyma develop prior to penetration of the host root. The sclerenchymatous cells form a characteristic structure, described as the sclerenchyma prong. During penetration the intrusive part of the haustorium reaches not only the host xylem but continues growing downwards until it entirely splits the host root. Comparable to a guillotine, the sclerenchyma prong is directly involved in this remarkable process. The sclerenchyma prong finally lies in the distal part of the haustorium. Following this mechanical slicing of the host root, tube-like cells of the intrusive part actively penetrate the host xylem in an axial direction.     

Water Relations of the Mistletoe Amyema fitzgeraldii and its Host Acacia acuminata

Water relations of the mistletoe Amyema fitzgeraldii and itshost Acacia acuminata were studied near Geraldton, Western Australia.Transpiration rates of host and parasite under unstressed winterconditions varied more than 300–fold between day and nightwhile leaf water potential gradients between the partners remainedwithin the range 0·4–0·6 MPa. Plots of transpirationagainst leaf water potential indicated closely similar fluidphase resistances in host and parasite during daytime but divergentbehaviour at night due to an apparently large increase in resistanceof the haustorial junction between the partners. Data for summerand winter studies across a full range of light intensitiesshowed the parasite to transpire, on average, 1·4 timesfaster and to exhibit noticeably lower water use efficienciesthan its host. Experiments following restorative changes atnight in leaf water potentials of host and parasite on detachedhost branches supplied through their cut ends with water indicatedthat the haustorium offered a major resistance to water uptakeby the parasite. Restoration of leaf water potentials by theparasite lagged markedly behind that of the host, especiallyduring winter, leading to a rapid build up in potential gradientbetween partners. A phase of rapid flow into the parasite thenfollowed, presumably motivated by lowering of the haustorialresistance. Reversal of the potential gradient between hostand parasite was recorded in a night-time study involving abagged (non-transpiring) mistletoe attached to a host branchfrozen at the base to prevent further water uptake. Mechanismsare proposed to account for the apparently highly variable natureof the resistance of the haustorium. Key words: Mistletoe, transpiration, haustorial resistance, Amyema fitzgeraldii, Acacia acuminata     

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     

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     

The Regulation of the Water Potential Gradient in the Host and Parasite Relationship betweenSorghum bicolorandStriga hermonthica

A glasshouse experiment was carried out to investigate the factorscontrolling the abstraction of xylem fluid from its host bythe parasiteStriga hermonthica(Scrophulariaceae).Strigahad amean daily transpiration rate far exceeding that of its hostsorghum (Sorghum bicolor), with infestation byStrigaalso shownto lower the transpiration rate of the host. Stopping the host'stranspiration was shown to decrease the transpiration rate ofthe parasite. Stopping the parasite's transpiration only gavean initial increase in the host's transpiration rate which wasnot sustained. The parasite had a lower water potential thanits host, values being -0.42 MPa and -0.23 MPa, respectively,and an accompanying higher osmotic pressure of 0.68 MPa against0.51 MPa for sorghum. Modifying the water potential gradientby bagging both partners together showed that the differentialin osmotic pressure and water potential was largely maintainedby the parasite's higher rate of transpiration. A favourablewater potential gradient towards the parasite still existedfollowing the cessation of transpiration, this being generatedby the haustorial resistance to hydraulic conductivity whichwas found to be some 1.5–4.5 times greater than that offeredby the parasite shoot. Both the high rate of transpiration andthe increased resistance across the haustoria would appear tobe necessary means to facilitate the diversion of host resourcesto the parasite.Copyright 1997 Annals of Botany Company Striga hermonthica; sorghum; water relations; haustorium; root parasite     

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     

Development of Tapinanthus bangwensis (Engler and Krause) Danser and Contact with the Host

Tapinanthus bangwensis forms a single large union with the host.Initially, the haustorium penetrates the host-cortex by thepressure of growth and enzymic action. On contact with the wood,the haustorium induces meristematic activity in the xylem parenchymaand cambium which leads to the dissection of the host wood andpenetration of haustorial branches between the dissected portions.Vascular contact is made by the formation of adjacent conductingcells in the host and parasite tissues.     

Xylem Transport and Storage of Amino Acids by S.W. Australian Mistletoes and their Hosts

Amino acid composition of xylem (tracheal) sap and ethanolicextracts of shoots of mistletoes (Amyema spp. and Lysiana casuarinae)and their hosts were compared, using material collected in theirnative habitats. Data indicated that certain host xylem soluteswere transferred directly to the parasite xylem, while otherswere either not absorbed or were metabolized prior to transfer.Certain solutes were major constituents of parasite xylem, butundetected or only in trace amount in the host. Shoot aminoacid pools of parasites differed markedly from those of hosts.The mistletoe, Amyema preissii, exhibited differential storageand transport of arginine when parasitizing three differentspecies, but accumulated proline on only two of these hosts.Host- specific amino acids (djenkolic acid in Acacia saligna,and tyramine in Acacia acuminata) were transported and accumulatedin relatively large amounts by the parasite, but were not detectedin other associations. Proline was the major solute of Amyemalinophyllum parasitizing Casuarina obesa, but arginine predominatedin Lysiana casuarinae on the same host. However, when L. csuarinaeparasitized A. linophyllum, in turn parasitic on C. obesa, theLysiana accumulated equal amounts of proline and arginine andmore asparagine than when directly on the Casuarina. Xylem feedingof ¹⁵N-labelled aspartic acid or ¹³N-(amide labelled) asparagineto cut shoots or whole haustoria-bearing plants of the mistletoeA. preissii resulted in 68–73% of the ¹⁵N of aspartateand 24–30% of that of asparagine appearing in ethanol-solubleshoot amino compounds other than the fed solute. ¹⁵N labellingpatterns of detached shoots were not noticeably different fromthat of whole plants suggesting that the haustorium had relativelylittle effect on processing incoming solutes. Alanine, glutamine,and arginine were principal recipients of ¹⁵N from aspartate,alanine and glutamine in the case of fed asparagine. It is estimatedthat 24% of the carbon requirements for dry matter accumulationin Amyema linophyllm were met by intake of xylem sap solutesfrom its host Casuarina obesa. Key words: Amino acids, xylem transport, mistletoes, host: parasite relations, N metabolism     

Influence of Flooding on Net CO2Assimilation, Growth and Stem Anatomy of Annona Species

A series of experiments was conducted to assess net CO₂assimilationand growth responses to waterlogging of grafted and seedlingtrees in the genus Annona. Seedlings of A. glabra, A. muricataandA. squamosa L., and scions of ‘Gefner’ atemoya(A. squamosaxA. cherimola Mill.), ‘49-11’ (‘Gefner’atemoyaxA. reticulata L.), ‘4-5’ (‘Priestley’atemoyaxA. reticulata), A. reticulata grafted onto either A.glabra, A. reticulata orA. squamosa rootstocks were floodedfor up to 60 d. Soil anaerobiosis occurred on the third dayof flooding. Seedlings ofA. glabra and A. muricata, and thescions ‘49-11’, ‘Gefner’ atemoya, andA. reticulata grafted onto A. glabra rootstock were consideredflood tolerant based on their ability to survive and grow inflooded conditions. Scions of the normally flood-sensitive A.reticulata, ‘Gefner’ atemoya, and ‘49-11’tolerated root waterlogging when grafted onto the flood-tolerantspecies, A. glabra. In contrast, flooding of A. squamosa seedlingsand rootstocks, and A. reticulata rootstocks greatly reducedgrowth and net CO₂assimilation rates, and resulted in 20–80%tree mortality. Stem anatomical responses to long-term flooding(12 continuous months) were assessed in seedlings of A. glabraand A. muricata, and trees of ‘49-11’ grafted ontoA. glabra. Flooded trees developed hypertrophied stem lenticels,particularly in A. glabra, and enlarged xylem cells resultingin thicker stems with reduced xylem density. Flooding did notincrease air spaces in pre-existing xylem near the pith or inxylem tissue that was formed during flooding. Thus, flood tolerancedid not involve aerenchyma formation in the stem. Copyright1999 Annals of Botany Company Flood tolerance, net CO₂assimilation, photosynthesis, stem anatomy, shoot growth, anaerobiosis, Annonaceae.     

Xylem Hydraulic Characteristics, Water Relations and Wood Anatomy of the Resurrection PlantMyrothamnus flabellifoliusWelw.

Myrothamnus flabellifoliusWelw. is a desiccation-tolerant (‘resurrection’)plant with a woody stem. Xylem vessels are narrow (14 µmmean diameter) and perforation plates are reticulate. This leadsto specific and leaf specific hydraulic conductivities thatare amongst the lowest recorded for angiosperms (k_s0.87 kg m^-1MPa^-1s^-1;k_l3.28x10^-5kg m^-1MPa^-1s^-1, stem diameter 3 mm). Hydraulic conductivitiesdecrease with increasing pressure gradient. Transpiration ratesin well watered plants were moderate to low, generating xylemwater potentials of -1 to -2 MPa. Acoustic emissions indicatedextensive cavitation events that were initiated at xylem waterpotentials of -2 to -3 MPa. The desiccation-tolerant natureof the tissue permits this species to survive this interruptionof the water supply. On rewatering the roots pressures thatwere developed were low (2.4 kPa). However capillary forceswere demonstrated to be adequate to account for the refillingof xylem vessels and re-establishment of hydraulic continuityeven when water was under a tension of -8 kPa. During dehydrationand rehydration cycles stems showed considerable shrinking andswelling. Unusual knob-like structures of unknown chemical compositionwere observed on the outer surface of xylem vessels. These maybe related to the ability of the stem to withstand the mechanicalstresses associated with this shrinkage and swelling.Copyright1998 Annals of Botany Company cavitation, desiccation, hydraulic conductivity, refilling, resurrection plant, root pressure, xylem anatomy,Myrothamnus flabellifolius