Freezing in Conifer Xylem: I. PRESSURE CHANGES AND GROWTH VELOCITY OF ICE

To determine whether freezing causes wide-spread cavitationin the xylem of freezing trees, pressure and temperature weremeasured inside freezing conifer sapwood blocks. Pressure risesof up to 3.3 MPa were recorded and average radial growth velocitiesof ice were between 1.75 and 2.3 µm s^–l. These growthvelocities of ice are less than the minimum growth velocityfor bubble nucleation during freezing. To complement this experimental study finite difference modelsof freezing in a single tracheid and freezing in an idealizedtree stem were constructed. The single tracheid model predictspressure rises similar to those measured experimentally. Thismodel also predicts that 5% to 8% of water in a tracheid lumenmigrates out of the tracheid during freezing. The tree stemmodel predicts growth velocities of ice three times faster thanthe values measured experimentally. These results are compared with previous contradictory theoriesof freezing in conifers. Key words: Freezing, xylem     

针叶树基因组特征及其序列资源挖掘进展

针叶树是裸子植物中最大也是分布最广的一支。作为多年生木本植物,针叶树不仅为工业提供建筑、造纸等重要原料和其它可再生能源,而且在北半球的生态平衡中也起着重要作用。因其独特的分类地位、重要的经济价值和生态价值,针叶树序列资源挖掘备受重视。然而其庞大且复杂的基因组阻碍了这一进程,截至2013年4月,尚无获得全基因组序列的针叶树种。随着第2代测序技术的出现及生物信息学的快速发展,针叶树序列资源挖掘也从转录组过渡到全基因组测序,后者己在松属（Pinus）、云杉属（Picea）和黄杉属（Pseudotsuga）部分树种中开展。该文首次归纳了针叶树基因组特征．回顾了针叶树序列资源挖掘进程,并重点介绍了火炬松（Pinustaeda）、欧洲云杉（Piceaabies）和白云杉（Piceaglauca）的全基因组测序项目。     

The Cohesion-Tension theory of sap ascent: current controversies

In recent years, the Cohesion-Tension (C-T) theory of sap ascentin plants has come under question because of work publishedby Professor Ulrich Zimmermann and colleagues at the Universityof Wrzburg, Germany. The purpose of this review is to (1) statethe essential and testable elements of the C-T theory, (2) summarizethe negative evidence for the C-T theory, and (3) review criticallythe positive evidence for the C-T theory and the evidence thatthe Scholander-Hammel pressure bomb measures xylem pressurepotential (P_x) correctly, because much of the evidence for theC-T theory depends on pressure bomb data. Much of the current evidence negates the conclusions drawn byZimmermann from studies using the xylem pressure probe (XPP),but it is not yet clear in every instance why the XPP resultsdisagree with those of other methods for estimating xylem pressure.There is no reason to reject the XPP as a useful new tool forstudying xylem tensions in the range of 0 to –0.6 MPa.Additional research is needed to test the C-T theory with boththe XPP and traditional methods. Key words: Cohesion-Tension theory, cavitation, embolism, xylem pressure probe, pressure bomb     

Some Constituents of Xylem Sap and their Possible Relationship to Xylem Differentiation

Bleeding sap of Actinidia chinensis and Betula populifolia andguttation fluidof Avena sativa were analysed for sugars, amino-acids,auxin, and certain enzymes. A wide range of amino-acids wasfound in all three. Auxin was not detected in the bleeding sap,but was present in Avena guttation fluid (5.1 µg IAA equivalent/l).‘IAA oxidase’, acid phosphatase, ribonuclease, deoxyribonuclease,and protease were detected in the bleeding sap and guttationfluid. The possibility that some of the substances found insap and guttation fluid are products of autolysing, differentiatingxylem cells in the roots is discussed.     

针叶树多倍体及其在遗传育种中的应用研究

多倍体及多倍化已成为生命科学研究的热点,林木中,众多速生高抗品种的成功选育和应用给林木遗传育种带来了新的研究热潮.作为主要的用材树种针叶树,天然多倍体极少,研究材料极其缺乏,严重限制了多倍体的应用研究.大量多倍体植株的获得以弥补其天然发生不足,是促进多倍体针叶树研究发展的前提.利用体细胞胚胎发生技术结合染色体工程技术等现代生物技术取代传统组织培养,解决了针叶树组织培养中生长缓慢、生根困难的瓶颈,是获得人工多倍体针叶树的最重要途径.     

Fixing and Staining of Conifer Tissues with Lacto-Propiono-Dimethylsulfoxide Carmine

During a study of the development of callus from immature microsporophylls and megagametophytes of Pinus spp. (Bonga 1974), tissue squashes were stained with aceto-carmine (Gerlach 1969). However, this stain had some drawbacks. The cells and nuclei formed in the pollen and pollen tubes during the early stages of culture were often obscured by large numbers of granules (cf. Christiansen 1973), and often, small groups of small vigorous cells within the callus failed to stain.     

Terpenoid Biosynthesis and Specialized Vascular Cells of Conifer Defense

Defense-related terpenoid biosynthesis in conifers is a dynamic process closely associated with specialized anatomical structures that allows conifers to cope with attack from many potential pests and pathogens. The constitutive and inducible terpenoid defense of conifers involves several hundred different monoterpenes, sesquiterpenes and diterpenes. Changing arrays of these many compounds are formed from the general isoprenoid pathway by activities of large gene families for two classes of enzymes, the ter...     

The Characteristics of the Adjustment in the pH of the Xylem Exudate of Segments of Vigna Hypocotyls during Xylem Perfusion

The xylem vessels of segments of Vigna hypocotyl were perfusedwith solutions of various pH values. With both acid and alkalineperfusion solutions, the pH of the xylem exudate tended to equilibrateat approximately 6 during the course of xylem perfusion. Thisphenomenon was observed both in air and under anoxia, and alsoin segments pretreated with methanol or preheated in a microwaveoven. Therefore, the adjustment in pH does not depend on therespiratory metabolism but originates in the buffering actionof the xylem wall apoplast. High concentrations of protons inducedthe release of other cations into the xylem exudate under anoxia,and high concentrations of K⁺ or Ca²⁺ ions induced the releaseof protons into the xylem under anoxia. These results indicatethat a cation-exchange reaction on the xylem cell wall was responsiblefor the buffering effect. The physiological role of the highbuffering capacity of the xylem wall apoplast is discussed inlight of the role of the proton pump of the plasmalemma in elongationgrowth. (Received September 25, 1990; Accepted January 21, 1991)     

On the Conductivity of the Xylem in Trees

The xylem of three tree species, the ring porous Fraxinus excelsiorL., and the diffuse porous Acer pseudoplatanus L., and Salixatrocinerea Brot., has been examined to determine how closelyflow through it obeys the Poiseuille equation. The equationwas most closly obeyed in the relationship between flow-rateand pressure difference between the ends of the xylem cylinder,though there were differences between the ring and diffuse poroustypes. The relationship of flow-rate and specific conductivityto the radius of the xylem cylinder was complex.Fraxinus showeda rise with increase of radius up to a given size (0.54cm),followed by a steep decline. This decline was shown to be causedby a progressive increase in the number of vessels which wereunable to conduct water. Small increases in the viscosity ofthe xylem liquid produced a disproportionately large fall inthe rate of flow, and the specific conductivity of the xylemdecreased. With viscosities greater than 2.5 centipoise, thespecific conductivity remained constant. An effect of alternatelyraising and lowering the hydrostatic pressure of the xylem wateron the specific conductivity of the wood is described. These results are discussed in relation to the structure ofthe xylem, and particularly with regard to the possibility thatthe conduction channels in the wood consist of a system of microcapillariessurrounding the vessel cavities.     

Sap Ascent in Vascular Plants: Challengers to the Cohesion Theory Ignore the Significance of Immature Xylem and the Recycling of Munch Water

In recent years the cohesion theory has been attacked on thegrounds that direct measurements made with the pressure probeindicate that sap tensions are much less (maximum tension approx.0.7 MPa) than indicated by parallel measurements made with themore conventional methods: osmotic methods, pressure bomb, orpsychrometer. It has also been claimed that other direct methodsdo not support the cohesion theory. Thus a re-examination usingthe Renner technique indicated sap tensions of approx. 2.5 MPa.Also an independent method based on mercury penetrometry providesevidence that sap tensions of at least 2.0 MPa can be demonstrateddirectly implying, that serious limitations arise from the pressureprobe method itself. Without tensions exceeding 2.0 MPa mangroveswould be unable to extract fresh water for transpiration fromseawater. It is suggested that the pressure probe is susceptibleto bias because it investigates the least mature xylem conduitswhile they are still under varying degrees of turgor pressureand only partially interconnected with the main xylem system.This supposition is supported by claims that the xylem sap sampledby the probe contains significant concentrations of solutes.Additionally water, supplied by reverse osmosis from the sievetubes (‘Münch water’), is continually beingliberated in the vicinity of the outermost xylem vessels hydratingthem to an atypical degree which can explain several of thediscrepancies claimed. These results, which are supported bythe work of others, demonstrate that the challenges to the cohesiontheory for the ascent of sap are ill-founded. The release ofwater from the phloem can explain not only some discrepanciesclaimed by the cohesion challengers, but also explain the refillingof cavitated xylem conduits: a hitherto unsuspected role forthe phloem transport system. Cohesion theory; sap ascent; cavitation; pressure probe; xylem transport; vessel development; recycled water; reverse osmosis     

Relationship of Xylem Embolism to Xylem Pressure Potential, Stomatal Closure, and Shoot Morphology in the Palm Rhapis excelsa

Xylem failure via gas embolism (cavitation) induced by water stress was investigated in the palm Rhapis excelsa (Thumb.) Henry. Xylem embolism in excised stems and petioles was detected using measurements of xylem flow resistance: a decrease in resistance after the removal of flow-impeding embolisms by a pressure treatment indicated their previous presence in the axis. Results supported the validity of the method because increased resistance in an axis corresponded with: (a) induction of embolism by dehydration, (b) increased numbers of cavitations as detected by acoustic means, (c) presence of bubbles in xylem vessels. The method was used to determine how Rhapis accommodates embolism; results suggested four ways. (a) Embolism was relatively rare because pressure potentials reach the embolism-inducing value of about −2.90 megapascals only during prolonged drought. (b) When embolism did occur in nature, it was confined to the relatively expendable leaf xylem; the stem xylem, which is critical for shoot survival, remained fully functional. (c) Even during prolonged drought, the extent of embolism is limited by complete stomatal closure, which occurred at the xylem pressure potential of −3.20 ± 0.18 megapascals. (d) Embolism is potentially reversible during prolonged rains, since embolisms dissolved within 5 h at a pressure potential of 0.00 megapascals (atmospheric), and xylem sap can approach this pressure during rain.     

Xylem Transport and the Negative Pressures Sustainable by Water

Experimental measurements of water's ability to sustain negativepressures are reviewed with special emphasis on the relevanceof the results to xylem transport. Results vary over severalorders of magnitude and depend on the conditions of testing.The greatest negative pressures are measured using heated, pressurizedwater or purified, degassed water. When conditions more closelyapproximating those found in biological systems are used, wateris considerably weaker. From these measurements, one can predictthe most likely range of negative pressures that can be sustainedin the xylem. This range is between -0·1 and -0·6MPa (absolute pressure). Negative pressures near -1 MPa arepossible, but require stringent conditions. Negative pressuresgreater than -2 MPa are also possible, but unlikely, based onthe experimental evidence.Copyright 1994, 1999 Academic Press Cavitation, embolism, negative pressure, xylem transport, Z-tube     

Conifer Woods of the Pliocene Age from Yunnan, China

The Tertiary floras play an integral role in understanding the biodiversity and interactions between climate and vegetation in Yunnan, China. The fossil spores, pollen grains, and leaves in this region have been investiagedintensively. In comparison, the woods have been studied relatively little. A large number of Pliocene wood specimens was collected from the opencast lignite coal mine of Hongxing situated in Changning County of Yunnan Province. Among the collection, Tsuga cf. dumosa (D. Don) Eichler and Pinus cf. armandii Franchet were identified based on wood structures. The subtle feature of tori extensions is reported in the wood of T. cf. dumosa. Considering the climatic requirements of modern T. dumosa and P.armandii, the two species of conifer described from Pliocene sediments probably grew in mountainous terrain at an elevation of approximately 2 300 m, in a cool and humid environment.     

A Comparison of Root- and Shoot-wood Development in Conifer Seedlings

Tracheid diameter, wall thickness, and wall area per tracheidincreased towards the root tip in first-year seedlings of Piceasitchensis and Pinus sylvestris; this is shown to be associatedwith differences in duration of the expansion and wall thickeningphases of tracheid development. Transfer to short days producesa similar response throughout the plant, tracheid diameter andwall thickness decreasing shortly after transfer, due to a declinein rate of development. These data are discussed in relationto growth-regulator gradients likely to exist between root andshoot. It is pointed out that though effects of growth regulatorson tracheid dimensions are well documented, it has yet to beestablished whether this is due to their regulation of the rateor of the duration of development.     

Conifer seedling distribution and survival in an alpine-treeline ecotone

The importance of seedling establishment to the position ofalpine-treeline is recognized, yet little is known about factorsaffecting the survival of seedlings of treeline conifers during their initialyears of growth and establishment. This establishment period may have thegreatest mortality of all life stages until death of mature trees by disease orfire. Spatial and temporal patterns in the distribution and survival ofseedlings of Picea engelmannii and Abieslasiocarpa were evaluated over four years in analpine-treeline ecotone of the Snowy Range, Wyoming, USA. Seedlings andsaplings of both species occurred most frequently near islands of adult trees.For P. englemannii, this appeared partly due to decreasedsurvivorship of young seedlings (< 5 cm height) with greaterdistance away from tree islands. Survival of emergents of P.engelmannii was 28% greater on the north compared to southsides of tree islands, 48% lower on south-facing slopes comparedto other aspects, and 70% greater with overhead cover such as treebranches. Survival of emergents was greater in microsites with grass cover(90% survival) compared to without ground cover (44% survival),but lowest in microsites surrounded, but not covered, by grass (19%).From 1994–1999, natural seedling emergence and survival washighest in 1995 (80% survival of 221 P.engelmannii,and 100% of seven A. lasiocarpa, in a sample areaof432 m²), when the smallest mean difference in dailymaximum and minimum temperatures occurred, and lowest in 1994 (30% ofseven P. engelmannii), when above-averagetemperatures were accompanied by low rainfall and clear skies. The growthseasons of 1994 and 1995 had among the lowest and highest precipitation of theprevious 30-year period, respectively. In an artificial seedingexperiment, less than 20% of seedlings of both species survived theirfirst complete year of growth. In the autumn of the second year, almost25% additional mortality was observed in the remaining experimentalseedlings when they were exposed to clear, cold skies without the normalprotection of snowcover. Both spatial and temporal patterns of seedlingsurvivalsuggest that exposure to high sunlight may exacerbate low-temperatureandwater stress in young conifer seedlings, inhibiting their establishment in thisalpine treeline.     

Transport of Minerals in tree Xylem

Stem segments of poplar devoid of leaves were supplied withlabelled phosphate solution under pressure through the cut base.Characteristic build-up curves of phosphate were obtained alongthe stems. The curves showed an initial stage of increasingrate of build-up, due to diffusion before the profile front,followed by a linear stage due to profile movement in vessels,and ending with a stage of decreasing build-up rate followedby a saturation plateau. It was concluded that the stage ofdecreasing build-up rate was due to restricted movement of isotopein partially blocked or recently differentiated vessels. Consequencesof this are discussed in relation to mass velocity measurementsin xylem and some comparisons are made with phloem movement.     

Xylem conduction and cavitation in Hevea brasiliensis

Clones of Hevea were studied in an attempt to discover the reasonsfor differences in the hydraulic performance of xylem. Differencesbetween clones were determined, including hydraulic conductivityand conduit width and length distributions. However, it hasproved difficult to reconcile anatomical differences with physiologicalperformance for use in future plant breeding programmes. When leaf relative water content (RWC) had been reduced fromabout 95% to 85%, the hydraulic conductivity of petioles decreasedsharply to about 40% of the initial value. This value correspondedwith xylem sap tensions of 1.8–2.0 MPa. Acoustic detectionexperiments revealed that this reduction in hydraulic conductivitycoincided with the greatest occurrence of cavitation. It seemsinescapable that the reduction in hydraulic conductivity wascaused by embolization; thereafter gas bubbles blocked the flowof water inside many of the conduits. There was some indicationthat eventually such bubbles might be dissolved, because thehydraulic conductivity increased again if specimens were fullyrehydrated. Apparently, the incidence of cavitation coincides with the entryof gas bubbles via ultramicroscopic pores into the conduitsthrough the walls according to the air-seeding hypothesis. Whena petiolate leaf is tested in a pressure chamber it is impossibleto make satisfactory measurements of a balancing pressure beyondc. 1.8–2.0 MPa, because air bubbles, mixed with sap andescaping from the conduits, form a persistent froth. Xylem transportin Hevea seems to be disrupted relatively easily under waterstress which is a feature of other tropical species adaptedto rainforest–type environments Key words: Hevea, xylem, cavitation, conduit, hydraulic conductivity