Seasonal Changes in Nitrogen Fractions of Coptis japonica, a Temperate Forest Evergreen Chamaephyte, and their Ecological Implications

Seasonal changes in several forms of nitrogen were investigatedin Coptis japonica, an evergreen rosette hemicryptophyte intemperate deciduous forest. The concentration of total nitrogenin rhizomes and roots decreased during the period of new shootgrowth from winter to spring. In the rhizomes, total solubleprotein stored by early summer decreased gradually until winter,coupled with an increase in free amino acids. Nitrogen was largelystored in free amino acids in the roots, especially during summer.The total soluble protein in current-year leaves decreased fromspring to summer and then increased during winter. The seasonalchanges in nitrogen components were coincident with the changein light-saturated photosynthetic rates recorded in a previousstudy. The ratio of total soluble protein to total nitrogendecreased from spring to summer and then increased from latesummer to winter in the current-year leaves. In contrast, chlorophyllcontent and the ratio of chlorophyll to total nitrogen werehigher in summer than in other seasons. The results indicatethat nitrogen was used in a manner that better utilizes thevery weak light in summer and the higher light intensities inother seasons. The major component of the free amino acid poolwas asparagine, in every organ throughout the season, exceptfor the senescent leaves. Since asparagine has a high N:C ratio,we suppose that the asparagine-dominated amino acid pool isadvantageous in the carbon-limited environment of the forestfloor.Copyright 1994, 1999 Academic Press Free amino acid composition, total nitrogen, total soluble protein, photosynthesis, evergreen hemicryptophyte     

Jurassic Petrified Filician Plants from the Rajmahal Hills, India

Three types of petrified filician rhizomes and their associatedpetioles and roots from the Rajmahal Hills, Bihar, India aredescribed. Two rhizomes possess protostelic vasculature withmesarch protoxylem elements and exarch protoxylem points. Theformer are gleicheniaceous while the latter is described asa schizaeaceous rhizome. The associated petioles possess typicalgleicheniaceous anatomy while the roots show variations in thestructure of epiblema and cortex. filician fossils, Jurassic, Actinostelopteris pakurense, Gleichenia sonajoriense, Solenostelopteris jurassica, Gleichenioamyelon diarcha, Filicoamyelon cryptogrammoides Filicoamyelon actinostachyoides     

Plant Growth and Survival under Strict Anaerobiosis

Anaerobic incubation of seedlings and rhizomes reveals interspecificdifferences in the ability of seedlings and rhizomes of higherplants to survive under prolonged and strict anaerobiosis. Rhizomesof several species were killed by an anaerobic incubation at22 °C for 7 d while others survived and showed normal shootextension on return to aerobic conditions. A third group ofspecies showed healthy and geotropically normal bud and shootextension while their rhizomes were in the oxygen free environment.A detailed comparison of bud and shoot growth rates was madewith Scirpus maritimus under aerobic and anaerobic conditionsover a 14 d period. Under similar conditions four species of grass seedlings weresubjected to anaerobic conditions for 2–8 d at 5°C,15 °C and 25 °C. Seedling mortality was highest at 25°C in all four grasses. Interspecific differences were evident.The wetland species were more tolerant of anoxia than the grassesfrom drier habitats.     

Nitrogen Fixation Associated with Rinsed Roots and Rhizomes of the Eelgrass Zostera marina

Nitrogen fixation was associated with the rinsed roots and rhizomes of the seagrass, Zostera marina L. Nitrogenase activity (acetylene reduction) was greater on rhizomes compared to roots, and on older roots and rhizomes relative to younger tissue. Compared to aerobic assays, anaerobic or microaerobic conditions enhanced the rate of acetylene reduction by rhizomes with attached roots, with the highest activity (100 nanomoles per gram dry weight per hour) occurring at pO₂ = 0.01 atmosphere. Addition of glucose, sucrose, or succinate also increased the rate of acetylene reduction under anaerobic conditions, with glucose providing the most stimulation. In one experiment, comparison of acetylene reduction assays with ¹⁵N₂ incorporation yielded a ratio of about 2.6:1. Seagrass communities are thought to be limited by the availability of nitrogen and, therefore, nitrogenase activity directly associated with their roots and rhizomes suggests the possibility of a N₂-fixing flora which may subsidize their nutritional demand for nitrogen.     

Catalase Activity and Post-anoxic Injury in Monocotyledonous Species

Three anoxia-intolerant species, Glyceria maxima, Juncus effususand Iris germanica (var. Quechei), and three anoxia-tolerantspecies Schoenoplectus lacustris, Acorus calamus and Iris pseudacoruswere chosen for investigation. Rhizomes of anoxia-intolerantspecies show increased catalase activities when returned toair after periods of prolonged anoxia. Levels of catalase remainedfairly constant in anoxia-tolerant species under the same conditions.In the anoxia intolerant G. maxima, the post-anoxic increasein catalase activity was reduced by circulating the anaerobicatmosphere. This treatment also reduced the ethanol contentof the tissue under incubation, and increased the survival ofthe rhizomes as seen in their ability to resume growth in thepost-anoxic phase. Exposure of anaerobic G. maxima rhizomesto ethanol vapour increased post-anoxic levels of catalase activityand when this produced a 5-fold increase always resulted indeath of the rhizomes. Acetaldehyde vapour applied in the sameway gave rise to increases in catalase activity followed byrapid death of the rhizomes. It is suggested that post-anoxic oxidation of anaerobicallyaccumulated ethanol may result in a surge of acetaldehyde production,which could exert a toxic effect on the recovering tissues.The possible role of catalase in an ethanol-oxidation reaction,which is well documented in animals, is discussed in the lightof the association between the natural accumulation of largeconcentrations of ethanol and subsequent post-anoxic death insome plant tissues. Key words: Catalase, post-anoxia, ethanol     

Change in Size and Cell Number during the Development of Lateral Root Primordia in Zea mays L.

Increase in cell number, anlage volume and length have beeninvestigated during the overall period of lateral root primordiumdevelopment in excised primaries and in attached roots of Zeamays L. Each of these aspects of anlage growth was found toincrease more or less exponentially during the interval betweenprimordium initiation and subsequent emergence as a lateralin both batches of roots. Values were then determined for celldoubling time (Td), the size of the proliferative fraction (Pf),and for anlage volume (Tv) and length (Tt) doubling times duringthe overall period of primordium development and at intervalsduring this period in both the excised and attached roots. Thepattern of change which took place in Td, Tv, Tl and Pf duringlateral primordium development was found to be similar in bothbatches of roots. However, the overall period of anlage developmentwas shorter in the excised roots than in the attached ones.Moreover, when laterals grew out of the excised roots they didso with fewer cells than comparable laterals emerging from theattached roots. Zea mays L., maize, root primordia, lateral emergence, cell doubling time     

Mechanisms Controlling Changes in Water Movement Through the Roots of Helianthus annuus L. During Continuous Exposure to Oxygen Deficiency

The effects of oxygen deficiency on suction-induced water flux(Jv) through detached roots of hydroponically-grown sunflowerwere investigated over a period of up to 6 d. Jv was reducedby the time the oxygen partial pressure in the solution, spargedwith oxygen-free N₂, had fallen below 2 kPa (air – 20.6kPa). This reduction resulted from a decline in the hydraulicconductivity of the radial pathway for water movement (Lp),although up to 46% of the decline was attributable to changesin the osmotic component of the driving force (). Lp (mm s^–1MPa^–1), for aerobic roots was 2.23 ? 10^–5 and foranaerobic roots during the initial 16 h, 1.21 ? 10^–5. At 22 h after the onset of anaerobic treatments, Jv and Lp increased,with Lp values becoming 3 times greater than those measuredbetween 4 h and 16 h of treatment and 1.4 times greater thanin aerated roots. However, Lp increased a further 15-fold whenroots were killed by immersion in boiling water for 2 min. Duringand up to 6 d of anaerobic treatment, some roots retained Lpvalues similar to those at 22 h, while others displayed characteristicstypical of dead roots. At no time was there any indication ofreduced axial conductivity due to xylem vessel blockage. The results are discussed in terms of possible energy sourcesfor the maintenance of root integrity and their importance toplant survival during long periods of severe oxygen shortage. Key words: Anaerobiosis, oxygen deficiency, hydraulic conductivity, osmotic potential, water     

Localization of Thermo-Osmotically Active Partitions in Young Leaves of Nuphar lutea

The submerged roots and rhizomes of the aquatic vascular macrophyteNuphar lutea (L.) Sm. are aerated, at least in part, by pressurizedventilation. Depending on temperature differences of up to 5K between the inside of young, just-emerged leaves and the surroundingair, pressure differences of 79 to 100 Pa higher than atmosphericare detectable inside the lacunuous spongy parenchyma of theleaf blades. The pressurization is a consequence of structuralfeatures of leaf tissues separating the air filled spaces ofthe spongy parenchyma from the atmosphere. These tissues areacting as thermo-osmotic partitions. Whereas the dimensionsof the stomatal openings (about 5·6 x 2·4 µm)and of the intercellular spaces of the palisade parenchyma (diametersabout 15 µm) are too large, those of the monolayers ofcells separating the palisade and the spongy parenchyma (diameters:0·7–1·2 µm) are small enough to impedefree gaseous diffusion. This inner non-homogeneous partitioninggives rise to the so-called Knudsen diffusion, a physical phenomenonleading to pressurization of the warmer air inside the spongyparenchyma. The rising pressure difference is strong enoughto establish an air flow through the aerenchyma of the wholeplant and out of the most porous older leaves in which a temperatureinduced pressurization is never detectable. These thermo-osmoticallyactive leaves enhance the influx of air to the rhizome and thediffusion path for oxygen to the roots is shortened to the distancebetween rhizome and root tips. Therefore, pressurized ventilationin Nuphar is seen to be of considerable ecological importancefor plant life in anaerobic environments. Key words: Aeration, leaf anatomy, thermo-osmosis of gases, Nuphar lutea     

Seasonal Dynamics of Nitrogenous Compounds in a Nitrophilic Weed II. The Role of Free Amino Acids and Proteins as Nitrogen Store in Urtica dioica

The free amino acid fraction, representing the most importantnitrogen store in the overwintering below ground plant partsof Urtica dioica, consisted mainly (up to 80%) of asparagineand arginine. While asparagine was dominant in rhizomes, upto one year old, argrnine was specifically accumulated in theolder rhizomes and roots. In spring first the nitrogen storedin asparagine and with a delay of about three weeks that inarginine was mobilized and translocated to the rapidly growingshoots. Proteins may function as nitrogen store too, in particularin the case of the seeds. In the below ground organs, however,a special storage protein, rich in amides and/or arginine, whichwould correspond to the large quantities of these amino acidsin the overwintering organs, could not be detected. (Received September 3, 1984; Accepted November 2, 1984)     

Translocation of Photoassimilates Between Sister Ramets in Two Rhizomatous Forest Herbs

Autoradiography and liquid scintillation techniques were usedto trace the pattern of photoassimilate translocation in twoperennial forest herbs, Aster acuminatus Michx. and Clintoniaborealis (Ait.) Raf. Vegetative shoots of C. borealis emergeearly each spring from the growing apices of a long-lived rhizomesystem. Vegetative shoots of A. acuminatus emerge in late springfrom rhizomes that decay within 2 years. In both species rametssurvive for only one growing season. Any connected plants aresisters. Mature leaves of these two species were exposed for1 h periods to ¹⁴CO₂ during spring and summer. Radioactivitycould subsequently be detected in exposed leaves (16–84per cent, depending on the season), adjacent above ground plantparts (0.7–23 per cent), roots and rhizomes (4.9–84per cent) and, when present, flowers (4–8.5 per cent).Old rhizomes of C. borealis are prominent storage sites forphotosynthate. In A. acuminatus, no significant translocationbetween sister ramets (i.e. above ground shoots connected bya common rhizome) was observed. In C. borealis, there was small,but consistent translocation between sister ramets (0.2–4per cent). Disturbance of unexposed sister ramets by defoliation,shading or herbivory increased the flow of photoassimilatesto disturbed parts in C. borealis, but not in A. acuminatus.Based on the absence of translocation flow, ramets of A. acuminatusmay be regarded as physiologically independent. Connected rametsof C. borealis show physiological integration. These resultsare correlated with ecological differences between the two species. Aster acuminatus, Clintonia borealis, translocation, ramet, vegetative reproduction, forest herb     

Changing Kinetic Properties of Barley Alcohol Dehydrogenase during Hypoxic Conditions

The Finnish barley cultivar (Hordeum vulgare L, cv. Hankkija-673)was grown in solution culture for periods of one and two months.The one month old plants had seminal roots, whereas the twomonth old plants grew adventitious roots from the lower nodes.The roots were tested for ADH activity during aeration, during3 d of hypoxic treatment (brought about by passing nitrogenthrough the nutrient solution), and during a 4 d recovery period.The ADH activity, calculated on a protein basis, rose about4-fold during the nitrogen treatment, the greatest increaseoccurring in the adventitious roots. Differences in the kineticproperties of ADH during the hypoxic period were also foundbetween seminal and adventitious roots. The K_m for acetaldehydechanged little in the seminal roots during the hypoxic treatment,but in the adventitious roots it decreased considerably. Thephysiological significance of these changes is discussed. Key words: Hordeum vulgare, alcohol dehydrogenase, hypoxia, enzyme kinetics     

Origin of Exogenous Adventitious Roots in Rhizomes of the Fern Microsorium scolopendria (Polypodiaceae)

Adventitious roots of rhizomes of the fern Microsorium scolopendriaarise exogenously from apical initials which have their originin the cell layer directly beneath the epidermis. Divisionsof the apical initial and its derivatives produce the adventitiousroot apical meristem. Cells between the apical cell and themeristele of the rhizome develop into the root trace. Anatomy, light microscopy, Microsorium scolopendria Burm.) Copel     

Light-dependent Anaerobic Induction of the Maize Glyceraldehyde-3-Phosphate Dehydrogenase 4 (GapC4) Promoter in Arabidopsis thaliana and Nicotiana tabacum

The maize glyceraldehyde-3-phosphate dehydrogenase 4 (GapC4)promoter confers strong and specific anaerobic gene expressionin tobacco (Nicotiana tabacum) and potato (Solanum tuberosum).Here we show that the promoter is also anaerobically inducedin Arabidopsis thaliana. Histochemical analysis demonstratesthat the promoter is anaerobically induced in roots, leaves,stems and flower organs. Surprisingly, the strong anaerobicinduction of the promoter is dependent on light and on the substitutionof oxygen with carbon dioxide. High carbon dioxide concentrationalone does not induce the promoter in the presence of oxygenand light. If anaerobic conditions are generated under completedarkness or if plants are submerged, no induction above backgroundis observed. When transgenic tobacco harbouring a GapC4 promoter–reportergene construct is analysed for light dependent anaerobic induction,the results are indistinguishable from those with arabidopsis.The implications for using the GapC4 promoter as an anaerobicreporter for monitoring alterations in the anaerobic signaltransduction pathway are discussed.     

Diurnal Rb+ Transport from Roots to the Laminar Pulvinus in Phaseolus vulgaris L.

The primary leaves of kidney bean (Phaseolus vulgaris L.) openunder light and close in the dark by the deformation of thepulvinus resulting from diurnal distribution changes of K⁺,Cl^–, organic acid (or H⁺) and NO₃^–. When Rb⁺ was added as a tracer of K⁺ to the seedlings throughtheir roots, it was transported to the pulvinus cells duringthe light period but not during the dark period. Transpirationoccurred vigorously in the light but almost stopped in the dark.We concluded that Rb⁺ absorbed by the roots was carried to thepulvinus by the transpiration stream. Phaseolus vulgaris L., pulvinus, Rb⁺, diurnal transport transpiration stream     

Some Effects of Indol-3-yl Acetic Acid on Lateral Root Development in Attached and Excised Roots of Pisum sativum L.

The effect of continuous exposure to indol-3-yl acetic acid(IAA) on primordium initiation and their subsequent emergenceas lateral roots was determined in excised and attached rootsof Pisum sativum. IAA was found to stimulate the number of primordiainitiated per centimetre of attached or excised primary. Similarly,lateral emergence in terms of the number produced per centimetreof primary was promoted in the presence of IAA. This stimulationof lateral emergence even took place in excised roots whichwere 1 cm in length at the onset of culture and which neverproduced secondary roots over a 6-d culture period when grownin the absence of auxin. These effects of IAA on lateral rootdevelopment have been considered in relation to the concurrentchanges which take place in proliferative activity in the apicalmeristem of the primary root during exposure to auxin. Pisum sativum, garden pea, anlage, primordium, emerged lateral, cell proliferation, indol-3-yl acetic acid     

Seasonal Dynamics of Nitrogenous Compounds in a Nitrophilic Weed I. Changes in Inorganic and Organic Nitrogen Fractions of the Different Plant Parts of Urtica dioica

The seasonal dynamics of nitrogenous compounds in the differentplant parts of Urtica diocia, a perennial nitrophilic weed,were investigated under natural conditions during the courseof a year. In the below ground plant parts (roots and rhizomes)a nitrogen store was built up during summer and autumn, consistingmainly of free amino acids. In the seeds, however, proteinswere of major importance as nitrogen reserve. In early springthe below ground nitrogen store was quickly mobilized, translocatedto the upper plant parts and used for the protein synthesisof the shoots, enabling them a rapid growth prior to the developmentof their own capacity for nitrogen assimilation. (Received September 3, 1984; Accepted November 2, 1984)     

The Importance of Water Vapour for the Circulating Air Flow through Nelumbo nucifera

Aquatic vascular plants depend on an adequate oxygen supplyin order to maintain growth and reproduction in anaerobic environments.Nelumbo nucifera is able to survive with a gas transport systemwhich supplies oxygen to the roots and rhizomes submerged inthe anaerobic sediment. It was possible to demonstrate thatthis gas transport system is based on a purely physical phenomenonThermo-osmotic oxygen transport was first demonstrated on freshleaves with the help of an oxygen-sensitive electrode. A definiteenhancement of oxygen flow was obtained through excised leaveswhen a temperature difference between the ambient and lacunarair was present in light. These leaves were then dried to brittlenessand the enhanced oxygen flow was still detectable. This showsthat not only photosynthetic oxygen, but also atmospheric oxygencan be transported to the buried organs. The absolute flow ofoxygen through dry leaves was much lower than through freshleaves, but the thermo-osmotic transport of oxygen still functioned.Furthermore, the process of thermo-osmosis need not rely ona difference in humidity between the two sides of a porous partition,but may be linked causally to the temperature difference andthe pore size. Key words: Nelumbo nucifera, oxygen transport, thermo-osmosis of gases     

A Study of Stelar Ultrastructure in the Heterosporous Water Fern Marsilea quadrifolia L

Sieve tube elements occur in the rhizomes and petioles of Marsileaquadrifolia. These are either thick walled with compound sieveplates in oblique end walls or thin walled with simple sieveplates in transverse end walls. Vessels are restricted to themetaxylem in the roots where the phloem contains sieve cellsonly. The sieve pores are invariably callose lined and as inother pteridophytes, excepting the Lycopsida, refractive spherulesare ubiquitous in the sieve elements of Marsilea. The luminaof the protoxylem tracheary elements in the rhizomes and petiolesare occluded by tyloses but probably remain functional in theroots. Pericycle cells backing on to the root protoxylem armspossess wall ingrowths. Transfer cells are however absent fromthe vascular tissue of the rhizomes and leaves. It is suggestedthat their presence in the root pericycle is related to theretrieval of ions from the xylem sap which may be particularlycritical in water plants. The incidence of transfer cells incryptogams appears to be far more sporadic than in angiosperms.The root endodermis of Marsilea possesses a casparian stripand abundant vacuolar tannin deposits. Plasmalemmasomes arenumerous adjacent to the pericycle transfer cells. vascular ultrastructure, Marsilea quadrifolia L, transfer cells, sieve tube elements, tyloses     

Assimilation and translocation of nitrogen and carbon in Curcuma alismatifolia Gagnep

Curcuma or Siam tulip (Curcuma alismatifolia Gagnep.) is an ornamental flowering plant with two underground storage organs, rhizomes and storage roots. Characteristics of N and C assimilation and transport in curcuma were investigated. The plants were treated with ¹⁵NH₄⁺ + ¹⁵NO₃^? and ¹³CO₂ at 10, 13 or 21 weeks after planting. Plants were sampled at several stages up to 32 weeks. The C stored in old storage roots was used rapidly during the first 10 weeks; after which N stored in old rhizomes and old storage roots were used. The daily gain in C depending on photosynthesis was remarkably high between 10 and 21 weeks. However, the daily gain in N was relatively constant throughout the growth period. The ¹⁵N absorbed at 10 weeks was initially accumulated in leaves and roots, but some was transported to flowering organs at 13 weeks. At harvest, 41% of ¹⁵N was recovered in new rhizomes and 17% in new storage roots. After ¹³CO₂ exposure at 10 and 13 weeks, the distribution of ¹³C among organs was relatively constant in subsequent stages. When given ¹³CO₂ at 21 weeks, a large amount of labelled C was recovered in new storage roots and new rhizomes at harvest. Both new rhizomes and new storage roots stored N and C, however, rhizomes played a more important role in supplying N, while storage roots provided C.     

Different patterns of aerenchyma formation in two hygrophytic species of Paspalum (Poaceae) as response to flooding

Paspalum modestum and P. wrightii are perennial grasses growing in permanent and seasonally flooded areas, respectively. The former produces short rhizomes and floating culms, the latter forms long rhizomes and erect culms. Variations in percentage aerenchymatous space (PAS) in different organs as a response to flooding was analysed using a clone of each species. Eighteen plantlets of each clone were cultivated during 7 months under flooded vs. unflooded conditions. After this period, roots, rhizomes, culms, and leaf sheaths were collected and prepared. PAS was measured using an image analysis device, and data were analysed using ANOVA.Production of aerenchyma took place in both species within the cortical parenchyma of roots, rhizomes and culms, and the mesophyll of leaf sheaths, both in flooded and unflooded plants. Under flooding conditions PAS increased in both species, although the individual response of organs differed: whereas in P. modestum PAS increased primarily in substratum-fixed roots, in P. wrightii all organs produced additional aerenchyma uniformly. Contrasting responses are understood as adaptations to permanent and seasonal flooding, respectively.