The Export and Distribution of 14C-labelled Assimilates from each Leaf on the Shoot of Lolium temulentum during Reproductive and Vegetative Growth

In both reproductive and vegetative plants of Lolium temulentumL., the export of ¹⁴C-labelled assimilates from each healthyleaf on the main shoot to terminal meristem, stem, tillers,and roots was measured each time a new leaf was expanded, fora period of 5 to 6 weeks. Some labelled assimilates moved fromeach leaf on the main shoot to every meristem in the same shoot,as well as to the tops and roots of adjacent organically attachedtillers. The terminal meristem of the reproductive shoot, which includedthe developing inflorescence, received 70–80 per centof the carbon assimilated by the emerged portion of the growingleaf, 15–25 per cent of the carbon assimilated by thetwo youngest expanded leaves, and 5–10 per cent of thatfrom each of the older leaves. A similar pattern of carbon supplyto the terminal meristem was found in vegetative shoots, exceptthat older leaves on young vegetative shoots supplied even lessof their carbon to the terminal meristem. The general conclusionis that developing leaves at the tip of the shoot receive aboutthe same proportion of carbon from each leaf as does a developinginflorescence. Young expanded leaves provided most labelled assimilates forstem growth; during both reproductive and vegetative growth,expanded leaves increased their export of labelled carbon tostem, and exported less of their ¹⁴C to roots and sometimesto tillers. In these reproductive and vegetative shoots, grown in a constantexternal environment, the major changes in the pattern of distributionof labelled assimilates appeared to be the result of increasedmeristematic activity in stem internodes; the development ofan inflorescence had no obvious direct effect on the carboneconomy of shoots.     

Patterns of 14C-labelled Assimilate Partitioning in Red and White Clover During Vegetative Growth

A quantitative analysis of the ¹⁴C-labelled assimilate suppliedby the expanded leaves on the primary shoot to growing leaves,stem, lateral shoots (branches or stolons) and roots in redand white clover was conducted during vegetative growth. Stem growth of the primary shoot was inhibited in both cloversand utilized no energy resources. The growing leaves at theprimary shoot apex of white clover imported 4 per cent of theshoot's assimilate compared with 10 per cent in red clover.At the basal end of the primary shoot, the tap root of whiteclover imported 16 per cent of the shoot's assimilate comparedwith 22 per cent in red clover. Branches in red clover and stolonsin white clover were by far the largest sinks for primary shootassimilate, importing 39 per cent and 63 per cent of the labelledassimilate, respectively. Analyses of the translocation of assimilate from individualprimary shoot leaves demonstrated that in both clovers olderleaves exported more of their assimilate to branches or stolons,whereas younger leaves exported more of their assimilate toroots, and possibly in white clover, to growing leaves at thetip of the shoot. Of the labelled assimilate exported to branchesor stolons, each primary shoot leaf exported preferentiallyto the branch or stolon in its own axil, but in addition exportedsubstantial quantities of assimilate to all other axillary shoots,particularly those arising from basal axils where the subtendingleaf had died. Trifolium repens, Trifolium pratense, red clover, white clover, assimilate partitioning, perennation     

Distribution patterns of assimilated 14C in vegetative and reproductive shoots of Lolium perenne and L. temulentum

The movement of ¹⁴C-labelled assimilate to the terminal meristem, stem, mature leaves, tillers and roots was measured in Loliurn perenn and Lolium temulentum after exposure to ¹⁴C0₂ of the youngest fully-expanded leaf and, on fewer occasions, the oldest healthy leaf on the main shoot. During early vegetative growth, the terminal meristem, tillers and roots received most of the ¹⁴C exported from the youngest leaf. As the shoot aged, more ¹⁴C was exported to the terminal meristem and tillers and less to roots. When the stem became a sizeable sink for ¹⁴C at the six-leaf (L. temulentum) or eleven-leaf (L. perenne) stage, less ¹⁴C moved to tillers and much less to roots. The terminal meristem continued to receive ¹⁴ at a steady rate throughout late vegetative growth. The transition from vegetative to reproductive growth in both species was marked by an abrupt increase in the export of ¹⁴C to stem from the upper leaf, but there was little change in the proportion of ¹⁴C which moved to the developing leaves and incipient inflorescence at the terminal meristem. At the same time, less ¹⁴C moved to tillers and much less to roots. Immediately before ear emergence, the export of ¹⁴C from the upper leaf (flag leaf) to the stem declined and the proportion moving to the ear increased, reaching a maximum of 55–75% as the ear emerged. The relative patterns of export of upper and lower leaves showed that while some ¹⁴ moved from each leaf to all meristems, the proximity of actively growing meristems appeared to be the main factor which determined the destination of most exported ¹⁴C. The distribution of ¹⁴C from upper and lower leaves was most alike in young vegetative plants of L. perenne. At later stages of development of both species, the terminal meristem and stem received most 14¹⁴C from the upper leaf, while roots and tillers received mos 14¹⁴C from the oldest leaf at the base of the shoot.     

Assimilate Distribution in Poa annua L.

The carbon economy of a flowering tiller of Poa annua L. hasbeen examined over the period from inflorescence emergence tograin shedding. The total import of ¹⁴C by the inflorescencereached a maximum at late grain filling but the relative importof assimilate was greatest 14 days after its appearance andrepresented 20–25 per cent of that assimilated by theinflorescence itself. The inflorescence continued to be an importantassimilatory organ after grain ripening when it exported morethan 50 per cent of its assimilate to the stem, roots and othertillers. The patterns of distribution of assimilates from the youngestuppermost and the oldest green leaf of the reproductive tillerwere largely determined by the stage of development of the inflorescence.The youngest leaf mainly supported the inflorescence up to theend of the grain-filling stage but then supplied assimilatesbasally to the roots and adjacent tillers. The oldest greenleaf supported the growth of the stem and the inflorescenceup to anthesis but after this supplied assimilates mainly tothe roots and tillers. Removal of grains or the entire inflorescence only 1 h beforesupplying ¹⁴CO₂ greatly reduced the rate of fixation of ¹⁴CO₂and the export of radiocarbon, as well as changing the patternof distribution of assimilates within the plant. The significanceof these results is discussed and comparisons made with cerealsand perennial grasses.     

Root-Tiller Interrelationships in Spring Barley (Hordeum distichum (L.) Lam.)

Root-tiller relations were investigated in spring barley grownin soil in deep pots. The total dry wt of the root system reachedits maximum 6 weeks from sowing, when the shoot weight was only50 per cent of its value at maturity. Seminal and nodal rootscomprised 40 and 60 per cent, respectively, of the total rootdry wt at maturity; the majority of the nodal root weight wasassociated with the main shoot. The main shoot had approximatelytwice as many nodal roots as either of the first two primarytillers (T1 and T2), and the primary and secondary tillers appearinglater were very poorly rooted. Some tillers, especially secondarytillers that died prematurely, produced no nodal roots. Theweight of the seminal roots and nodal roots attached to themain shoot continued to increase up to maturity but the drywt of nodal roots on tillers declined with time. This patternof growth was closely related to the pattern of ¹⁴C assimilateddistribution within the root system. A very small proportionof ¹⁴C assimilated by the main shoot and T1 and T2 was exported.The majority of the exported assimilate went to the seminalroot system and to nodal roots attached to the main shoot. Individualnodal and seminal roots seemed to have different roles in supplyingnutrients to the shoot system, with the former mainly providing³²P-phosphate to its tiller of origin and the latter generallysupplying the main shoot and primary tillers. Hordeum distichum. (L.) Lam., barley, root growth, nodal roots, seminal roots, tillering, assimilate distribution, ³²P-distribution     

Effect of Assimilate Supply on Development and Growth Potential of Axillary Buds in Roses

The effect of assimilate supply on axillary bud developmentand subsequent shoot growth was investigated in roses. Differencesin assimilate supply were imposed by differential defoliation.Fresh and dry mass of axillary buds increased with increasedassimilate supply. The growth potential of buds was studiedeither by pruning the parent shoot above the bud, by graftingthe bud or by culturing the bud in vitro. Time until bud breakwas not clearly affected by assimilate supply during bud development,Increase in assimilate supply slightly increased the numberof leaves and leaf primordia in the bud; the number of leavespreceding the flower on the shoot grown from the axillary budsubstantially increased. No difference was found in the numberof leaves preceding the flower on shoots grown from buds attachedto the parent shoot and those from buds grafted on a cutting,indicating that at the moment of release from inhibition thebud meristem became determined to produce a specific numberof leaves and to develop into a flower. Assimilate supply duringaxillary bud development increased the number of pith cells,but the final size of the pith in the subsequent shoot was largelydetermined by cell enlargement, which was dependent on assimilatesupply during shoot growth. Shoot growth after release frominhibition was affected by assimilate supply during axillarybud development only when buds sprouted attached to the parentshoot, indicating that shoot growth is, to a major extent, dependenton the assimilate supply available while growth is taking place.Copyright1994, 1999 Academic Press Assimilate supply, axillary bud, cell number, cell size, defoliation, development, growth potential, meristem programming, pith, Rosa hybrida, rose, shoot growth     

Partitioning of 14C Labelled Assimilates in Arctium tomentosum

Plants of the biennial Arctium tomentosum were grown from seedto seed-set in an open field under three different treatments:control plants receiving full light intensity, plants with aleaf area reduced by 45 per cent, and shaded plants receivingonly 20 per cent of natural illumination. At various stagesof development the youngest fully expanded leaf of one plantin each treatment was exposed to ¹⁴CO₂ for half an hour. Subsequentdistribution of labelled assimilates in various plant partswas determined after eight hours. In the first year, the mostdominant sink was the tap root irrespective of variation inassimilate supply. During the production of new vegetative growthin the second season, a larger amount of radioactive photosynthatewas recovered from above ground parts, especially during formationof lateral branches. Seed filling consumed 80–90 per centof labelled carbon exported from the exposed leaf. In the secondyear, the most pronounced difference between treatments wasin the degree of apical dominance, being highest in shaded plantsand lowest in the plants with cut leaves. Results from ¹⁴C experimentsagreed fairly well with a ‘partitioning coefficient’derived from a growth analysis of plants grown independentlyunder the same experimental conditions. Reasons for discrepanciesbetween the ¹⁴C results and the partitioning coefficient arediscussed. Arctium tomentosum, burdock, variation in assimilate supply, assimilate distribution, ¹⁴CO₂, labelling, growth analysis     

Long-term Partitioning, Storage and Re-mobilization of 14C Assimilated by Lolium perenne (cv. Melle)

The youngest fully expanded leaves of single tillers of vegetativeperennial ryegrass plants were exposed to ¹⁴CO₂. Thereafter,quantitative and fractional analysis of the partitioning, storageand re-mobilization after defoliation of the ¹⁴C-labelled assimilatewas sequentially conducted over a 22 d period. In undefoliated plants, most ¹⁴C reached its final destinationwithin 5–6 of feeding. Forty per cent of assimilated ¹⁴Cwas subsequently lost through respiration, while 13.5, 8.5 and34 per cent remained in roots, stem bases and tops respectively.At least some ¹⁴C was distributed to tillers throughout theplant, but secondary tillers subtended by the fed tiller madethe greatest demand on ¹⁴C translocated from the fed tiller. A small, but significant portion of ¹⁴C was invested into longterm storage in undefoliated plants, four per cent of the totalassimilated still being present in a labile chemical form inroots and stem bases 22 d after feeding. In plants that wereseverely defoliated 4 d after feeding, depletion of reserve¹⁴C was observed relative to undefoliated plants. The depletiontook place from stem bases, not roots, and both low and highmolecular weight storage compounds were involved. A portionof the depleted ¹⁴C was incorporated into new growth after defoliation. Lolium perenne, perennial ryegrass, assimilate partitioning, storage, re-mobilization, defoliation     

Assimilation and Translocation in Perennial Grasses

Single leaves, ears, or shoots of timothy (Phleum pratense L.)were exposed in light to ¹⁴CO₂, then left overnight, after whichthe plants were autoradiographed. The following conclusionswere drawn. Actively growing leaves retain all their assimilatesand import from older ones. Fully expanded leaves export butdo not import assimilates. Export begins before leaf expansionis complete, so import and export may for a time be simultaneous.Exports go at first to younger leaves and to roots, accumulatingat meristems. At later stages, exports move downwards ratherthan upwards. Buds and small tillers import from older shoots,but large tillers do not import from other shoots or exportto other large ones. Ears assimilate while still green, andimport assimilates from their associated flag leaves. Exportsfrom other leaves on flowering stems move downwards. These findings agree in general with those from other plants:they are discussed in relation to the vascular system of thegrass plant, and the need for further studies, particularlyquantitative ones, is emphasized.     

The Distribution of Assimilates in Lolium multiflorum Lam. Following Differential Defoliation

In seedling plants of Lolium multiflorum Lam. the tillers weredefoliated but the main shoot was left intact. Radiocarbon as¹⁴CO₂ was supplied to this shoot at different times followingtiller defoliation and the pattern of distribution of labelledassimilates was determined quantitatively. It was found thata greater proportion (approximately 10–20 per cent) ofexported assimilate was translocated to the cut tillers butalthough the proportion supplied to the root system was lessthe total radiocarbon incorporated by the roots was unchanged.This was brought about by a large increase in the export ofradiocarbon fixed by the intact shoot—up to 100 per centfollowing one treatment. These alterations in the organizationof the defoliated plant lead to a greater efficiency in thecarbon economy and are discussed in relation to the stress imposedby defoliation.     

The Interdependence of Tillers in Lolium multiflorum Lam.--a Quantitative Assessment

Experiments were made to determine the extent of reciprocaltransfer of products (derived from the assimilation of ¹⁴CO₂)between various parts of the young vegetative grass plant (Loliummultiflorum Lam.). When individual laminae on different tillerswere supplied with ¹⁴CO₂, 47–64 per cent of the fixedcarbon was exported after 24 h. The principal sinks were theroot system and the shoot or tiller to which the fed leaf wasattached. Other tillers also received significant quantitiesof radiocarbon. When whole tillers were supplied with ¹⁴CO₂the percentage of fixed radiocarbon exported within 24 h rangedfrom 14–31 per cent. Of this, 50–74 per cent wasrecovered from the root system (except in the case of exportfrom the youngest tiller) but exchange of material between tillersalso occurred.A reciprocity diagram is presented and it is concludedthat despite the magnitude of the exchange no tiller showedan over-all net gain or loss. The main shoot and the tillersdiffered in the extent of their carbon exchange and in theirdegree of independence. The oldest daughter tiller of the mainshoot was the most independent and the main shoot most interdependent.     

Studies in the Nutrition of Apple Rootstocks: II. Effects of Concentration of Iron in the Nutrient Solution on Growth and Chlorophyll Content

Rooted one-year shoots were grown for one season by sprayingtheir roots with nutrient solution. Iron supplied as Fe-EDTAat four concentrations resulted in plants which were respectively(a) severely chlorotic, (b) mildly chlorotic, (c) dark greenand healthy (controls), and (d) dark green but with slight reductionin growth. Severely deficient plants showed 40–70 per cent reductionsin growth as measured by fresh weight, shoot length, diameterincrease, leaf area, net assimilation and relative growth-rates.Dry weights were reduced 70–80 per cent and of the totaldry-weight increment a greater proportion remained in the leaves,which had a lower dry weight and higher water content per unitarea. However, because the initial old stem formed a greaterproportion of the total dry weight, the leaf area ratio remainedabout 11 per cent lower than in the controls. Severely deficientplants had, per unit of chlorophyll, a higher dry-weight increaseand net assimilation rate than the controls. Mild deficiency caused 10–20 per cent reductions in growthand net assimilation rate; the leaf area ratio was normal. Possible mechanisms of the effects of low iron supply are discussed,while the small growth reduction at the highest Fe-EDTA concentrationis attributed to chelate toxicity     

Effect of Rate of Photosynthesis on the Pattern of Assimilate Distribution in the Graminaceous Plant

Infrared gas analysis and a quantitative radiocarbon tracertechnique were used to measure photosynthesis and the distributionof ¹⁴C-labelled assimilate in Lolium temulentum and a uniculmbarley exposed continuously or intermittently to contrastinglight intensities. Plants grown for 10–20 d in low light(<50 W m^–2 of visible light) exported a greater proportionof their assimilate to growing leaves at the terminal meristemand a smaller proportion to their roots and tillers than equivalentplants in high light (150 W m^–2). A single day's exposure(8.4 h photoperiod) to a contrasting light regime elicited achange in the pattern of assimilate distribution in the samediurnal period, but 2–3 d exposure was required for asubstantial shift in the pattern of supply of assimilate tomeristems. The data indicated that in terms of assimilate distributioncomplete adaption to a new light regime is attained in about7 d.     

Effect of Gibberellic Acid on the Distribution of Products of Photosynthesis in Sunflower

Single leaves on growing sunflower plants were allowed to assimilate¹⁴CO₂. Gibberellic acid was applied to the same leaf or to theterminal bud or the roots, and the distribution of assimilated¹⁴C was determined at intervals of 1–96 h. Gibberellicacid had no significant effect on initial distribution of ¹⁴Cduring the period of rapid export from the leaf, but enhancedre-export from the roots after translocation from the leaf hadvirtually ceased. Most of the ¹⁴C exported from the roots accumulatedin the shoot tip. The site of application of the hormone wasof relatively minor importance. Wherever it was applied themajor effect was enhancement of movement from the roots to theshoot tip. Application to the terminal bud was most effectivein this respect. There was no evidence that gibberellic aciddirectly affected the transport system, but the data supportthe hypothesis that it increases the strength of the sink inthe shoot tip. Helianthus annuus L., sunflower, assimilate transport, gibberellic acid, phloem transport     

In Vitro Culture and Plant Regeneration in Vicia faba subsp. Equina (var. Spring Blaze)

Explants of stem, leaves, roots, and cotyledons from etiolatedaxenically grown Vicia faba seedlings were cultured on a rangeof media. Shoot organogenesis was only obtained with nodal stemand cotyledonary node explants when cultured on MS medium with3% sucrose, 2.0 mg 1^–1 BAP and 02 mg 1^–1 NAA. Callusproliferation accompanied shoot organogenesis from nodal stemexplants. Successive subculture of nodal stem callus resultedin proliferation of regenerative callus which contained severalshoot bud initials. The capacity for shoot regeneration fromthis callus was maintained for 9 months. Histological studiesreveal de novo formation of meristematic centres in callus andtheir further development into bud primordia. High frequencyrooting of these adventitious shoots was obtained on half-strengthMS medium with 1.5% sucrose, 0.1 mg 1^–1 NAA and 0.5 mg1^–1 kinetin. Key words: Vicia faba, adventitious shoots, axillary shoots, de novomeristem formation, organogenesis, tissue culture     

Patterns of Assimilate Distribution and Source--sink Relationships in the Young Reproductive Tomato Plant (Lycopersicon esculentum Mill.)

Patterns of distribution of ¹⁴C were determined in 47-day-oldtomato plants (Lycopersicon esculentum Mill.) 24 h after theapplication of [¹⁴C]sucrose to individual source leaves fromleaves 1–10 (leaf 1 being the first leaf produced abovethe cotyledons). The first inflorescence of these plants wasbetween the ‘buds visible’ and the ‘firstanthesis’ stages of development. The predominant sink organs in these plants were the root system,the stem, the developing first inflorescence and the shoot ‘apex’(all tissues above node 10). The contribution made by individualsource leaves to the assimilate reaching these organs dependedupon the vertical position of the leaf on the main-stem axisand upon its position with respect to the phyllotactic arrangementof the leaves about this axis. The root system received assimilateprincipally from leaf 5 and higher leaves, and the stem apexfrom the four lowest leaves. The developing first inflorescencereceived assimilates mainly from leaves in the two orthostichiesadjacent to the radial position of the inflorescence on thevertical axis of the plant; these included leaves which weremajor contributors of ¹⁴C to the root system (leaves 6 and 8)and to the shoot apex (leaves 1 and 3). This pattern of distributionof assimilate may explain why root-restriction treatments andremoval of young leaves at the shoot apex can reduce the extentof flower bud abortion in the first inflorescence under conditionsof reduced photoassimilate availability. Lycopersicon esculentum Mill, tomato, assimilate distribution, source-sink relationships     

Developmental anatomy of the reproductive shoot in <Emphasis Type="Italic">Hydrobryum japonicum</Emphasis> (Podostemaceae)

Podostemaceae are unusual aquatic angiosperms adapting to extreme habitats, i.e., rapids and waterfalls, and have unique morphologies. We investigated the developmental anatomy of reproductive shoots scattered on crustose roots of Hydrobryum japonicum by scanning electron microscopy and using semi-thin serial sections. Two developmental patterns were observed: bracts arise either continuously from an area of meristematic cells that has produced leaves, or within differentiated root ground tissue beneath, and internal to, leaf base scars after an interruption. In both patterns, the bract primordia arise endogenously at the base of youngest bracts in the absence of shoot apical meristem, involving vacuolated-cell detachment to each bract separately. The different transition patterns of reproductive shoot development may be caused by different stages of parental vegetative shoots. The floral meristem arises between the two youngest bracts, and is similarly accompanied by cell degeneration. In contrast, the floral organs, including the spathella, arise exogenously from the meristem. Bract development, like vegetative leaf development, is unique to this podostemad, while floral-organ development is conserved.     

Photosynthesizing Leaves and Nodulated Roots as Donors of Carbon to Protein of the Shoot of the Field Pea (Pisum arvense L.)

In Pisum arvense, the amides and amino-acids normally suppliedto the shoot in the transpiration stream transfer carbon toprotein largely throught the amino-acids, aspartic acid (+asparagine),glutarnic acid (+glutamine), threonine, lysine, arginine, andproline. Carbon from carbon dioxide enters the protein of photosynthesizingtissues through an essentially complementary set of amino-acidsincluding glycine, alanine, serine, valine, and the aromaticamino-acids tyrosine, phenylalnine, and histidine. Young tissuesof the shoot synthesize certain amino-acids de novo by metabolismof sugars supplied from photosynthesizing leaves. Each mature leaf on a shoot contributes carbon to current synthesisof protein at the shoot apex. Sucrose accounts for more than90 per cent of the labelled carbon leaving any age of leaf whichhas been fed with ¹⁴CO₂. Upper leaves supply labelled assimilatesdirectly to the shoot apex, and the radiocarbon from these assimilatesis subsequently incorporated into a wide range of amino-acidunits of protein. The majority of the labelled assimilates exportedfrom a lower leaf move downwards to the root and nodules and,in consequence, the amino-acids and amides associated with rootmetabolism are strongly represented among the compounds eventuallylabelled in the apical region of the shoot.     

Significance of the simultaneous growth of vegetative and reproductive organs in the prostrate annual Chamaesyce maculata (L.) Small (Euphorbiaceae)

To elucidate the significance of the simultaneous growth of vegetative and reproductive organs in the prostrate annual Chamaesyce maculata (L.) Small (Euphorbiaceae) from the standpoint of meristem allocation, we investigated plant architecture, meristem allocation, and the spatial and temporal patterns in vegetative growth and reproduction in the reproductive stage. The numbers of secondary and tertiary shoots successively increased by branching in the reproductive stage, and the sum of shoot length was greater in secondary shoots than in primary shoots. The specific shoot length (shoot length per shoot biomass) was greater in lateral shoots than in primary shoots, indicating efficient lateral shoot elongation. The internode length was shorter in secondary shoots than in primary shoots, increasing the number of nodes per shoot length in secondary shoots. Many nodes on a shoot generated two meristems, one of which committed to a flower and one to a lateral shoot. The number of reproductive meristems was greatest in tertiary shoots, and 96% of total reproductive meristems on shoots were generated in lateral shoots. On almost all nodes, the reproductive meristem developed into a flower, and 95–98% of the flowers produced a fruit. Therefore, vegetative growth by branching in the reproductive stage contributed to the increase in reproductive outputs. From the standpoint of meristem allocation, the simultaneous growth of vegetative and reproductive organs in prostrate plant species might be important for increasing the number of growth and reproductive meristems, resulting in the increase in reproductive outputs.     

The shoot apex of Podostemaceae: De novo structure or reduction of the conventional type?

The paper reviews and discusses various interpretations of the shoot apex of Podostemaceae with special reference to subfamily Podostemoideae. Main questions concern (1) the proposed absence of a shoot apical meristem (SAM) in apical “meristemless” shoot tips of Podostemoideae and, as the consequence, the endogenous inception of leaf-borne leaves and branches and (2) the predicted stem bifurcation below a “terminal” dithecous (double-sheathed) leaf positioned instead of a shoot apex, as it is reported for subfamily Podostemoideae. Does the “meristemless” shoot apex represent a true evolutionary novelty? Does the view of stem bifurcation represent a new ramification pattern with the consequence that the “classical root–shoot model” of angiosperms is not valid for Podostemaceae? Both interpretations do not conform to previous studies that are complemented here by new data on the SAM of Zeylanidium olivaceum and Thelethylax minutiflora (Podostemoideae). Although a SAM is difficult to observe in the vegetative shoots of many Podostemoideae, it becomes well visible when the shoot passes into the flowering stage approaching the conspicuous shoot apex of floriferous shoots. The arguments of the absence of a SAM in vegetative shoots are not convincing and the endogenous origin of “leaf-borne leaves” appears questionable. Consequently, the “meristemless” shoot apex cannot be considered as a structure having evolved de novo. In the less advanced subfamilies Tristichoideae and Weddellinoideae, the leaf primordia develop only from a few apical cells of the outer shoot layer. This allows the conclusion that the surface layer of the apex in these subfamilies corresponds to the horizontally spread single-layered apical meristem of subfamily Podostemoideae. Similarly, the view of shoot bifurcation does not conform to the diachsial–sympodial branching pattern occurring in the cymose inflorescences of many Podostemoideae. This fact contradicts the presence of a terminal leaf.