光同化产物在裙带菜（Undaria pinnatifida）中的运转

本文报告了用１４Ｃ标记的光同化产物在裙带菜［Ｕｎｄａｒｉａｐｉｎｎａｔｉｆｉｄａ（Ｈａｒｙ．）Ｓｕｒ］孢子体中的运转现象。观察到光同化产物约需２０ｍｉｎ才能从叶片表皮进入中肋的髓部。在自然条件下,光同化产物主要自叶片梢部经中肋向生长部运转,在叶片梢部和生长部之间存在明显的源一库关系。运转速度约为４２～４８ｃｍ／ｈ。用组织放射自显影的方法证实了髓部的喇叭丝是主要的运转组织。向下运转的溶于酒精的光同化产物中,甘露醇占５０％以上。还在光同化产物中观察到游离的谷氨酸、天门冬氨酸和丙氨酸。     

THE RATE OF TRANSLOCATION IN ALARIA ESCULENTA (LAMINARIALES,PHAEOPHYCEAE)12

The rate of translocation of organic carbon in blades of Alaria esculenta (L.) Grev. (Laminariales) was calculated from blade growth data. The cross sectional area of sieve filaments in the midrib medulla was estimated from light microscopic examination of fresh material. Files of these filaments form a perimedullar ring occupying ca. 2/3 of the medulla. Values computed for specific mass transfer of carbon into the blade meristem ranged from 36.2 to 60.8 mg C.wk^?1.0.1 mm^?2 sieve filaments.     

PHYSIOLOGICAL INVESTIGATIONS ON ALARIA ESCULENTA (LAMINARIALES,PHAEOPHYCEAE). II. ROLE OF TRANSLOCATION IN BLADE GROWTH1,2

Laboratory studies on blade growth in Alaria esculenta (L.) Grev. showed 3 periods of rapid blade elongation during the year: October–November, February–April and late June. The first two periods are characteristic of many Laminariales; the unique June peak may reflect local nutrient conditions. While the distal blade functions as a source, supplying organic matter to the blade meristem, the stipe can be a source during periods of rapid growth or a sink during late summer when blade growth is slow. Maximum enhancement of elongation rate of blade meristems was observed in 40–50 cm blades; longer blades showed no further increase in growth rate. This blade length-growth promotion relationship may be independent of seasonal variations in meristematic activity. ¹⁴C tracer experiments suggested that separate growth promotion effects by distal blade, sporophylls and stipe were not additive in the intact thallus. The preferential source of assimilate for blade meristem growth was the distal blade. Secondary sources: sporophylls, which were activated following excision of the primary source; and stipe, which began to translocate assimilate when both sources were removed. The role of secondary sources in nature is discussed. Profiles of radioactivity in alcohol-soluble organic matter in blades are evaluated in relation to tracer profiles in higher plants and mechanisms of translocation.     

MORPHOLOGY AND TAXONOMY OF HIMANTOTHALLUS (INCLUDING PHAEOGLOSSUM AND PHYLLOGIGAS), AN ANTARCTIC MEMBER OF THE DESMARESTIALES (PHAEOPHYCEAE)1

The sporophyte of Himantothallus develops according to a closed pattern in which the number and position of the blades is determined by the location of trichothallic meristems in a filamentous germling. Expansion of the miniature juvenile to the massive adult thallus is accomplished by diffuse secondary growth and involves a change from filamentous rhizoids to a hapteroid holdfast, flattening of the stipe, and enormous increases in length, breadth, and thickness of both stipe and blade. The axis usually bears 1–8 lateral blades, often paired, and terminates in a flattened stub. Phaeoglossum is interpreted as a growth form of Himantothallus in which a terminal blade develops to the exclusion of lateral blades, the latter being represented by a single spine. Phyllogigas clearly falls within the morphological spectrum of Himantothallus, the lack of twisting being related to physical factors in the environment. Sporangia, interspersed with an equal or somewhat larger number of two-celled paraphyses, are borne in slightly elevated sori scattered over both surfaces of the blade. Zoospore germination was not observed, nor were gametophytes, either in culture or in the field. Haptera apparently originate from the meristoderm in the lower part of the maturing stipe and lack a filamentous medulla. The mature stipe and the mature blade are anatomically similar, being composed of a superficial meristoderm, a cortex of parenchyma-like cells, and a filamentous medulla. The meristoderm is usually a single layer of plastid-containing cells that divide anticlinally to accommodate (or effect) expansion and periclinally to produce cortical tissue inward. Cortical cells are in radial files and increase in diameter towards the interior. They usually are densely packed with physodes. The medulla is uniquely distinguished by the presence of sheathed trumpet hyphae. Cells of the trumpet hyphae have perforate end walls with callose deposits and probably function in conduction as do the sieve filaments in Laminariales. Sheathing cells are filled with plastids. Sheathing filaments form connections among themselves and with nearby unsheathed filaments. The sheathed trumpet hyphae and their matrix of unsheathed filaments form a plexus, which in the mature blade is flattened and may be stripped intact from the other tissues. Development of the embryonic sporophyte is very similar to that in Desmarestia, as is the anatomy of the adult thallus and the sporangia. From these considerations, Himantothallus is assigned to the Desmarestiaceae (Desmarestiales).     

Long Distance Transport in Macrocystis integrifolia: II. Tracer Experiments with C and P

Discs from mature regions of Macrocystis blades picked up significantly more [³²P]phosphate from the ambient medium than similar discs from young meristematic regions, and this uptake was higher in light than in darkness. Double-labeling experiments with NaH¹⁴CO₃ and [³²P]phosphate, using intact fronds as well as cut frond segments, indicated that ³²P was translocated from mature blades to sink regions at velocities of 25 to 45 centimeters per hour, velocities comparable to ¹⁴C translocation velocity in the same material. There was a slight delay in transport of ³²P which may be due to a delay in loading or to a high metabolism of ³²P in the transporting channels. Histoautoradiography of stipe segments in the translocation pathway indicated that transport of label occurred in the peripheral parts of medulla. An analysis of ³²P-labeled compounds in the fed blade and in the sieve tube sap, collected from basal cut ends of stipes, indicated major differences in labeling patterns. In the blade, a high proportion of ³²P was recovered as inorganic phosphate and relatively small amounts were found in hexose mono- and diphosphates, UDPG and ATP. In the sieve tube sap, however, only a small amount of ³²P was present as inorganic phosphate, a large proportion was found in hexose mono- and diphosphates, and appreciable amounts were present in ATP and UDPG.     

GROWTH OF BLADES OF NEREOCYSTIS LUETKEANA (PHAEOPHYTA) IN DARKNESS 1

Short term measurements were made of the relative growth rate of the fast growing portions of blades of Ncreocystis luetkeana (Merl.) Post. et Rupr. during exposure to natural daylight and during prolonged darkness. Growth was only slightly, but significantly, faster during the 12 h of daylight than during the 12 h that included 8 h of darkness. Clearly considerable growth occurred at night. In blades amputated 1 cm beyond the zone measured, growth was slower during both night and day. In continuous darkness growth continued for up to 12 days. It was not influenced by amputation of distal blade tissue but it was increased by severance from the bulb and stipe. A mean volume increase of 50% of the blade tissue was recorded. While the organic content decreased, the drop was half that required to support the increase in volume. There was some evidence against translocation. It is possible that cellular biochemical rearrangements allow a light-independent increase in organic material.     

Changes in 32P-phosphorus compounds during translocation in Laminaria digitata (L.) lamouroux

³²P was applied to a Laminaria digitata thallus and the pattern of ³²P phosphorylated compounds was studied, as a function of time, in the different tissues involved in translocation, i.e. source, pathway and sinks. The results showed that, 3 hours after absorption by the uptake region (lamina), the bulk of the radioactivity was incorporated into organic compounds (70 to 80% of total ³²P taken up), hexose monophosphates being the heaviest labelled. Further change in that region was marked by an accumulation of ³²P in the inorganic pool (65 to 70% after 13 days). Conversely, the ³²P pattern in the medulla of the stipe, which initially showed a similar pattern to the uptake region, did not vary during translocation. The pattern of ³²P distribution into sinks (growing stipe peripheral tissue or hapteron) leads to accumulation of the radioactive element in inorganic and acid-insoluble fractions. These results are discussed in terms of comparative distribution of ³²P in the different parts of the thallus and suggest that phosphate moves as Pi in that alga.Abbreviations TCA trichloroacetic acid - Po organic phosphate - Po sol acid-soluble organic phosphate fraction - Po insol acidinsoluble organic phosphate fraction - Pi morganic phosphate fraction - P lip lipidic phosphate - Np protein nitrogen - ATP adenosine triphosphate - ADP adenosine diphosphate - PEP phosphoenolpyruvic acid - PGA phosphoglyceric acid - G-1-P glucose-1-phosphate - G-6-P glucose-6-phosphate - UDPG uridine diphosphoglucose     

Effect of Nitrogen Deficiency upon Translocation of C in Sugarcane

Withholding nitrogen decreased the percentages of nitrogen and chlorophyll in the blades; reduced the total fixation of radioactive carbon dioxide at 15, 37, and 178 seconds; and changed the relative composition of fixation products. Translocation of radioactive photosynthate from the fed part down the attached blade and into the stalk was less in the plants deprived of nitrogen than in the control plants supplied with nitrogen. Both the percentage of total activity translocated and the velocity of transport were decreased by nitrogen deficiency. During a translocation period of 90 minutes the minus nitrogen blade retained more ¹⁴C-sucrose than the control in the fed part and the blade below the fed part, but it sent less ¹⁴C-sucrose to the sheath of the fed leaf. Thus translocation decreased with nitrogen deficiency not for lack of sucrose but for some other reason. Although withholding nitrogen decreased translocation of labeled carbon in and from attached blades, there was no effect upon transport in detached blades. The effect of nitrogen deficiency upon translocation may be indirect and secondary to the effect upon growth of the plant as a whole.     

Kinetic characteristics of photoassimilate translocation in Alaria esculenta (Laminariales,Phaeophyceae)

The kinetics of translocation of ¹⁴C-labeled photoassimilate were studied in the kelp, Alaria esculenta (L.) Grev., using a Geiger-Müller detector-probe to measure radioactivity in the source and sink regions of dumbbell-shaped explants cut from blades. Rapid tracer efflux from the source occurred for 4 days following a pulse of [¹⁴C]bicarbonate, with 40–60% of the initial activity remaining in the source after 10–14 days. Portions of source and sink tissue were analysed for distribution of radioactivity in mannitol, amino-acid, organic-acid and insoluble fractions. About 75% of the radioactivity in both source and sink at the end of the experiments was in soluble organic matter. The translocation velocity of the moving solute front (1.0-1.6 cm·h^-1) was derived from time-course profiles of tracer arriving in Alaria sinks. Relative rates of translocation, calculated from these profiles, yielded skewed curves, with maximum rates of import by the sink occurring 72–96 h after the source was pulsed.     

FIELD STUDIES ON THE GIANT KELP NEREOCYSTIS1,2

An intertidal population of Nereocystis luetkeana was investigated in the field, and techniques for measuring sporophyte growth were developed. Primary regions of growth in plants prior to reproductive maturity were located in the bases of the blades, upper stipe, and holdfast. After reproductive areas on blades appeared, growth was confined to 2 major regions: the blade bases and the junction between lower stipe and holdfast where new holdfast haptera were produced. Complete deblading halted growth at whatever stage the plants had attained at the time of the operation but did not kill the plants. Partial deblading resulted in low but measurable growth rates in the blade bases. The data presented suggest that the function of the blades is twofold: they bear reproductive structures in the mature plants and. they may provide, materials used, in rapidly growing regions.     

Sugar transport in conducting elements of sugar beet leaves

Autoradiography was used to determine the distribution of labeled sugar in conducting elements of the blade and petiole of sugar beet leaves at intervals ranging from 5 sec to 24 hr. The processes of assimilation by the green cells, collection of sugar in the minor veins and export in phloem elements were demonstrated visually. It appears that in minor veins sugar is translocated in companion cells rather than sieve tubes. In major veins translocation occurs in sieve tubes.     

TRANSLOCATION OF 14C IN MACROCYSTIS INTEGRIFOLIA (PHAEOPHYCEAE)1,2

Translocation patterns in the giant kelp, Macrocystis integrifolia Bory, were investigated in situ using ¹⁴C tracer; sources and sinks were identified. Export was first detected after 4 h of labeling; experiments were routinely 24 h continuous ¹⁴C application. Mature blades exported ¹⁴C to young blades on the same frond and on younger fronds, as well as to sporophylls and frond initials at the bases of the fronds. Blades <0.3 m from the apex imported and did not export; this distance did not change seasonally. In spring export from blades 0.3–1.25 m from the apex was exclusively upwards; older blades also exported downwards. In fall downward export began 0.5 m from the apex, and blades >2 m from the apex exported exclusively downwards. Carbon imported by frond initials, young fronds, and sporophylls in fall may partly be stored for growth in early spring. No translocation was seen in very young plants until one blade (secondary frond initial) bad been freed from the apical blade; this blade exported to the apical blade for a time, but imported when it began to develop into a frond. The second and third formed blades on the primary fronds (sporophylls also exported when <0.3 m from the apex, and later stopped. Frond initials and sporophylls on later-formed fronds did not export at all. The translocation pattern in M. integrifolia differs from that previously reported in M. pyrifera in seasonal change and in distances from the apex at which the changes take place.     

CONTRASTING PATTERNS OF ALLOMETRY AND REALIZED PLASTICITY IN THE SISTER SPECIES MAZZAELLA SPLENDENS AND MAZZAELLA LINEARIS (RHODOPOHYTA)1

Phenotypic differences between the low wave exposure Mazzaella splendens (Setchell et Gardner) Hommersand and the high exposure Mazzaella linearis (Setchell et Gardner) Fredericq could be due to plasticity or genetic differentiation. Common gardens were used to assess their levels of plasticity after describing allometric relationships. As thalli lengthened, stipe development for M. splendens almost ceased even though blades continued to expand, but M. linearis formed a larger stipe before developing a blade that continued to stay narrow at longer thallus lengths. Common gardens demonstrated that M. splendens regrown in the site of M. linearis produced a wider blade than M. splendens regrown in its natural low energy site and that M. linearis regrown in low wave energy either could not form a wider blade or became narrower than thalli from its high energy site. Tetrasporophytes of M. splendens produced a longer and thicker stipe in the high energy site, but the larger M. linearis stipe was not realized because its wider blades made it vulnerable to hydrodynamic removal. Mazzaella splendens therefore had low survivorship in the high wave energy site, and survivors were not long enough to reproduce. Survivorship and reproduction of M. linearis was similar in both environments. Some of the M. splendens and M. linearis characters are plastic, but this plasticity was insufficient for convergence of phenotypes, and blade width plasticity was maladaptive at least for M. splendens. Developmental systems producing the stipe and blade phenotypes of each species have undergone genetic differentiation.     

Translocation of assimilates in Undaria and its cultivation in China

Undaria cultivation on a commercial scale began in China only in the last decade. Today, Undaria pinnatifida is the main species under cultivation concentrated in two provinces, Liaoning and Shandong. The annual production in the early nineties was 8000-13 000 tons dry weight, which is two or three times the pre-1980 figures. The raft cultivation method maintaining the alga at the desired depths generally ensures the light saturated rate of photosynthesis on clear days, and enhances production. Under the cultivated condition, the calculated annual primary productivity of this alga is 160 gC m⁻² y⁻¹. Translocation of ¹⁴C-labelled photoassimilates in rapidly growing sporophyte of Undaria pinnatifida was studied in the open sea. Samples from different parts of the blade with counterparts exposed to tracer (NaH¹⁴CO₃) showed that the translocation that occurred mainly from the tip of the blade to the growing region had obvious source-sink relationship. It took 20 minutes to translocate the labelled photoassimilates from the epidermis, via cortex, to the medulla of the midrib, where rates of translocation averaging 42–48 cm h⁻¹ were observed in the open sea. Production experiments of tip-cutting of the blades showed an increased production of 9%. This revised version was published online in August 2006 with corrections to the Cover Date.     

Source-sink relations in maize mutants with starch-deficient endosperms

Partitioning and translocation of photosynthates were compared between a nonmutant genotype (Oh 43) of corn (Zea mays L.) and two starch-deficient endosperm mutants, shruken-2 (sh2) and brittle-1 (bt1), with similar genetic backgrounds. Steady-state levels of ¹⁴CO₂ were supplied to source leaf blades for 2-hour periods, followed by separation and identification of ¹⁴C-assimilates in the leaf, kernel, and along the translocation path. An average of 14.1% of the total ¹⁴C assimilated was translocated to normal kernels, versus 0.9% in sh2 kernels and 2.6% in btl kernels. Over 98% of the kernel ¹⁴C was in free sugars, and further analysis of nonmutant kernels showed 46% of this label in glucose and fructose. Source leaves of mutant plants exported significantly less total photosynthate (24.0% and 36.3% in sh2 and bt1 compared to 48.0% in the normal plants) and accumulated greater portions of label in the insoluble (starch) fraction. Mutant plants also showed lower percentages of photosynthate in the leaf blade and sheath below the exposed blade area. The starch-deficient endosperm mutants influence the partitioning and translocation of photosynthates and provide a valuable tool for the study of source-sink relations.     

Long Distance Transport in Macrocystis integrifolia: I. Translocation of C-labeled Assimilates

Long distance transport of ¹⁴C-labeled photoassimilate was studied in Macrocystis integrifolia Bory. Movement of label followed the source-sink relationship; mature blades closer to the holdfast with young 2° and 3° fronds transported mostly to the base, those closer to the frond apex transported mostly to the apex, and those in intermediate positions transported both acropetally and basipetally. The velocity of movement of ¹⁴C as computed both from study of intact fronds and exudate was in the range of 35 to 72 centimeters per hour and these estimates are on the low side. The composition of the translocate as determined from intact fronds was the same as that determined from exudate analysis; furthermore, this composition was nearly identical with that of the photosynthate (40 to 50% mannitol and 40 to 50% amino acids). From these data we conclude that the exudate represents the sieve tube sap and that there is little if any selectivity exercised in the loading and translocation of photoassimilate. An analysis of translocated label in the growing apex is presented and indicates that the synthesis of polymeric compounds such as laminaran, alginate, cellulose, lipids, and “protein” occurs in situ from the transported mannitol and amino acids. Detailed data on chemical composition of sieve tube sap from M. integrifolia and M. pyrifera (L.) C.A. Agardh are given and compared with the sieve tube sap from higher plants. Finally, we show that stipe segments, 60 to 100 centimeters long with three to six attached blades, are useful for translocation studies in Macrocystis.     

Translocation of C in the sugarcane plant during the day and night

The time-course of translocation of ¹⁴C from the blades of the sugarcane plant was investigated by analysis and radioactive counting of successive samples punched from a single blade. In 1 experiment, the time-course was studied by determining the specific activity of the carbon dioxide respired by the roots.

The rate of translocation, expressed as percentage, was highest immediately after the application of the radioactive carbon dioxide. Morning-made photosynthate translocated a higher percentage during the morning than during the afternoon in 90-minute periods in the light. Afternoon-made photosynthate translocated as well or better than morning-made photosynthate for the first hour in the light.

The leaf-disk data and the specific activity of the carbon dioxide respired by the roots corresponded by showing lower rates of translocation by night than by day for several successive days. Also, the translocation of ¹²C sucrose was slower at night.

The ¹⁴C sucrose translocated by day was made primarily by photosynthesis; the sucrose translocated by night was made primarily by the conversion of other labeled compounds, e.g. organic acids, organic phosphates, and insoluble residue.

The radioactive constituent of the residue, which was converted to sucrose, was tentatively identified as a glucose-xylose-glucuronic acid hemicellulose, with most or all of the ¹⁴C in the glucose moiety.

Translocation of sucrose may be triggered by different mechanisms during the night than the day. The conversion of insoluble residue to sucrose by increasing the osmotic potential at the source would favor a pressure-flow mechanism for nocturnal translocation; whereas translocation by day is thought to be a process of phototranslocation, a photoactivation of the translocation mechanism.

     

The microscopy of P-protein filaments in freeze-etched sieve pores

Intact vascular bundles from Nymphoides peltata (S.G. Gmel.) O. Kuntze, shown to have translocated carbon-14, were freeze-fractured and etched for electron microscopy. The interpretation of freezefractured and etched sieve pores and P-protein filaments seen in them is discussed. The entire widths of most of the sieve pores seen contained filaments separated by less than 100 nm. Their arrangement indicates too high a resistance to flow for pressure flow alone to drive translocation at known rates; pumps would be necessary at places along sieve tubes. However, calculations are presented to show that during the time taken to fix pores, by fast freezing or chemically, the filaments in them could rearrange and move further by Brownian and other motion than the distances between filaments which we need to measure. These calculations show that it is not possible, by microscopy alone, to answer the outstanding question “How are filaments arranged in translocating sieve pores?” with enough certainty to tell us whether pressure flow is adequate to explain translocation where filaments are present. The calculations are relevant also to microscopy of other cell structures which may move.     

Sink to source transition in developing leaf blades of tall fescue

This study aims to characterize the translocation of photosynthates within and from developing tall fescue ( Festuca arundinacea ) leaves at the time of transition from sink to source. The developing leaf contains a source, the exposed tip, and a sink, the growing basal portion. When the exposed tip of the developing blade is labelled with ¹⁴CO₂, it exports photosynthates exclusively to sinks within the developing blade until the blade reaches 80% of its final length, when photosynthates begin to be exported from the blade and pass through the collar to reach the growing sheath and the next expanding leaf. Concurrently, the previous mature leaves reduce their level of photosynthate export to the developing blade; export stops as soon as the sheath of the developing leaf elongates beyond 10 mm. Export from the mature leaves to the growing sheath and to the next expanding leaf blade increases rapidly. Thus, in a developing tall fescue leaf blade photosynthate importation and exportation are exclusive events: the expanding blade imports photosynthate from mature leaves until it reaches 80% of its final length, then exportation begins and importation ceases.     

Proline Accumulation in Water-stressed Barley Leaves in Relation to Translocation and the Nitrogen Budget

Mobilization of N from leaves of barley (Hordeum vulgare L.) during water stress, and the role of proline as a mobilized species, were examined in plants at the three-leaf stage. The plants responded to water stress by withdrawing about 25% of the total reduced N from the leaf blades via phloem translocation. Most of this N loss was during the first 2 days while translocation of ¹⁴C-photosynthate out of the stressed blade still remained active. Free proline accumulation in the blade was initially slow, and became more rapid during the 2nd day of stress. Although a major free amino acid, proline accounted for only about 5% of the total N (soluble + insoluble) retained in severely stressed blades. When the translocation pathway in water-stressed leaves was interrupted just below the blade by a heat girdle, a cold jacket, or by blade excision, N loss from the blade was prevented and proline began to accumulate rapidly on 1st day of stress. Little free proline accumulated in the blades until after the ability to translocate was lost. Proline was, however, probably not a major species of N translocated during stress, because proline N accumulation in heat-girdled stressed leaves was five times slower than the rate of total N export from intact blades.