VASCULAR PATTERNS IN STEMS OF ARACEAE: SUBFAMILY POTHOIDEAE

The stem vasculature of representative species of ten of the 11 genera in the Pothoideae was analyzed with the aid of films of series of transverse sections. In all species a leaf trace typically diverges from a continuing axial bundle before departing to a leaf, with the possible exception of Heteropsis. However, within this common organizational scheme a considerable range of variation exists, e.g., with respect to degree of branching of axial bundles, and distance from a leaf-trace branch to the point of leaf-trace departure to a leaf. In addition, we have found a wide variety of patterns of bud-trace organization in different genera of the Pothoideae. For example, prominent arcs composed of numerous bud traces occur in the central cylinder of Pothos, Pothoidium, and Heteropsis. Comparative anatomy leads to the conclusion that Pothos and Pothoidium more closely resemble Heteropsis than Anadendrum. Anadendrum should be dissociated from the tribe Pothoeae. With respect to other genera in the Pothoideae, our preliminary results suggest that each of the genera Anthurium, Culcasia, Zamioculcas, Acorus, and Gymnostuchys are highly distinct from each other.     

VASCULAR PATTERNS IN STEMS OF ARACEAE: SUBFAMILIES COLOCASIOIDEAE,AROIDEAE AND PISTIOIDEAE

The stem vasculature of ten genera of Colocasioideae and three genera of Aroideae was analyzed by films of series of cross sections. The technique was unsuited for the numerous tuberous genera of Aroideae (and Pistia), which have shortened internodes. The Colocasioideae has long been recognized as one of the most natural large assemblages in the Araceae, a concept further supported by information from stem anatomy and vasculature. All species examined have amphivasal axial bundles that undergo frequent anastomosis and bifurcation of a seemingly irregular kind. Syngonium is the only viny genus and is exceptional in a number of anatomical features which are associated with its unusual morphology. One of the principle points of diversity in the Colocasioideae is the presence or absence of a permanent cortical vascular system. All four genera with a permanent cortical system (Caladiopsis, Caladium, Xanthosoma, and Syngonium) are neotropical. In the Aroideae (and Pistioideae) all of the tuberous genera have a highly condensed vascular system. Genera with elongated internodes (Stylochiton, Lagenandra, Cryptocoryne) also have a similar pattern, which makes taxonomic comparisons based on stem vasculature in the Aroideae of little value. Branch trace insertion is much less well developed in Colocasioideae and Aroideae than in most other subfamilies.     

PATTERNS OF ENDOTHECIAL WALL THICKENINGS IN ARACEAE: SUBFAMILIES POTHOIDEAE AND MONSTEROIDEAE

A survey of the patterns of endothecial wall thickenings in 106 representative species from 20 genera in the Pothoideae and Monsteroideae was made using cleared anthers, sections and macerations. The wide variety of wall thickenings that is present is based on an annular-helical pattern. Variations in thickenings are related to differences in cell shape, cell orientation, intergradation between helical and annular patterns, pitch of helices, presence of branched thickenings, and various types of discontinuities in thickenings. Notable exceptions to the annular-helical pattern include Culcasia, which lacks a differentiated endothecial layer with thickenings, and Acorus, which has a peculiar stellate pattern that is unique in the family. No single pattern consistently characterizes either subfamily, although continuous helices are common in the Monsteroideae, and rare in the endothecium of Pothoideae (except Anadendrum). Monsteroideae frequently exhibit a series of slanted separate thickenings on anticlinal walls, which is absent from Pothoideae except in Heteropsis. The slanted pattern is considered a variation on a rectangular helix, involving discontinuities of thickenings on the periclinal walls. Some monsteroid genera show considerably more interspecific variation (Rhaphidophora) than others (Monstera). Endothecial thickenings constitute an anatomical character that is useful in the systematic study of Araceae; present results support other anatomical studies in identifying Culcasia and Acorus as highly divergent genera in the Pothoideae.     

SYSTEMATIC OCCURRENCE OF STEROLS IN LATEX OF ARACEAE: SUBFAMILY COLOCASIOIDEAE

Latex from representative New and Old World colocasioid species was analyzed by combined gas chromatography-mass spectrometry for the presence of sterols. Free sterols were detected in latex of all New World species examined, and were localized in ca. 1 μm terpenoid particles which are abundant and account for the white-opaque appearance of the latex. The sterol profiles were qualitatively and/or quantitatively distinct for each of the New World species examined. No free sterols were detected in latex of Old World genera examined, which are consistently clear to cloudy, and lack terpenoid particles. The presence of free sterols as small particles is typical of New World Colocasioideae with white latex. The results of this study provide chemical evidence for distinguishing Old and New World groups of Colocasioideae, which was recently proposed based on stem vasculature, pollen, and leaf type. This is the first report of sterol-rich latex in a monocotyledon.     

PATTERNS OF STAMEN VASCULATURE IN THE ARACEAE

A survey of the three-dimensional organization of stamen vasculature in 100 genera and over 350 species of Araceae was made using clearings. The Araceae exhibit highly varied stamen vasculature, with three main patterns: 1) vascular bundles unbranched, 1–3 per stamen, 2) forked bundles in some or all stamens, 3) anastomosing vascular systems with several to many bundles entering a single stamen. Three major groups of taxa in the family can be recognized on the basis of their predominant pattern of stamen vasculature. Virtually all genera with bisexual flowers (most Pothoideae, Monsteroideae, Calloideae, Lasieae) have unbranched bundles, one per stamen, except two to three in some species of Holochlamys, Spathiphyllum, and Scindapsus. Forked stamen bundles are virtually restricted to and occur nearly throughout the monoecious Lasioideae, Philodendroideae, Colocasioideae and among certain Aroideae (sensu Engler), including tribes Arophyteae, Spathicarpeae (Asterostigmateae) and Protareae. No forked bundles were found in tribe Areae (Aroideae), except Theriophonum indicum or any Araceae with bisexual flowers, except two species of Cyrtosperma. Anastomosing systems are virtually limited to members of tribe Areae with larger stamens, such as Arum, Helicodiceros, Eminium and Dracunculus species. A similar pattern occurs in some Amorphophallus, but other patterns occur as well. The distributions of forked bundles and anastomosing systems in the family are notable because they are both highly congruent with Philodendroideae-Colocasioideae, and Aroideae, respectively, in Grayum's new system for the family. Virtually all of the genera with forked bundles are grouped together in the Philodendroideae-Colocasioideae. All of the genera with anastomosing systems are in the Areae, including the complex and variable Amorphophallus, which has an uncertain systematic placement.     

POLAR TRANSPORT OF GROWTH-REGULATORS IN PITH AND VASCULAR TISSUES OF COLEUS STEMS

Movement of IAA-C¹⁴ and 2,4-D-C¹⁴ through cylinders of known size and histology was compared using liquid scintillation counting. Both auxins showed strongly polar movement, even through pith parenchyma cut from Coleus internode #5, the youngest internode to have ceased elongation. The polar movement was correlated with sizable elongation of the excised cylinders. Velocities of basipetal movement for a given auxin, as determined by the intercept method, showed small or negligible differences between pith and “corner” cylinders. (Corner cylinders comprised mostly vascular tissue, plus some cortical, pith, and epidermal cells.) For IAA, basipetal velocities ranged from 2.1 to 3.3 mm per hr; for 2,4-D, they were 0.6–0.8. For both auxins there was much more net loss into corner than into pith cylinders, a difference associated with the fact that corner cylinders showed 10 times as many cells in transection. More 2,4-D moved basipetally through corner than through pith cylinders and the reverse was true of IAA. By chromatographic evidence, all the radioactivity in the basal receiving blocks was still associated with the auxin molecules.     

SURVEY OF SHOOT ORGANIZATION IN THE ARACEAE

Shoot organization is examined in 87 species from 29 genera representing all six subfamilies of the Araceae and of Acorus, which has been placed in a separate family. Within each taxonomic group examined, the details of shoot organization are presented, including the types of segments and articles which make up the shoot, the degree of expansion of leaf blades, and the placement of buds along the shoots. Literature on shoot organization of the 29 genera is reviewed. The degree of correlation between shoot organization characteristics and systematic groupings is examined, and the utility of these characteristics for systematics is evaluated. It is found that within the taxa observed, the pattern of shoot organization provides a distinctive “fingerprint” at the generic or sectional level, sufficient for determination of the group. Some patterns which appear are pointed out: taxa with bisexual flowers usually produce a single inflorescence at the terminus of a vegetative article. A few taxa with bisexual flowers produce pairs of inflorescences at the ends of articles. Multiple inflorescences (more than two) at an article terminus occur only among taxa with unisexual flowers. Multiple inflorescences are associated with anisophyllous or homeophyllous sympodial growth, while single or paired inflorescences are associated with homeophyllous or intermittent homeophyllous sympodial growth. These patterns might be understood as the result of selection for flexibility of reproductive effort and of seasonal reproduction.     

A SURVEY OF FLORAL ANATOMY IN ARACEAE

Flowers of 23 species representing six subfamilies of Araceae were studied by means of serial cross sections, special attention being given to vascular patterns and to taxa of supposed phylogenetic importance. Floral structure is shown to be extremely diverse with no unifying pattern common to all subfamilies. Conclusions include the following: (1) Lysichiton has a specialized gynoecial vascular pattern which differs from others encountered in the survey and which weighs against the primitive position attributed to this genus by Hutchinson. (2) Philodendron, with its multiple stylar canals, cannot have originated from subfamily Pothoideae, as Engler's phylogenetic concept would require of all Araceae; instead, it appears that several syncarpous evolutionary lines have evolved independently from extinct apocarpous members of the family. (3) In Acorus, stamens are introrse and dorsal carpellary bundles are lacking; these characters and others justify the recognition of Acorus as a separate subfamily Acoroideae. In addition, the survey revealed a peculiar deterioration of the inner ovary wall and the septa in several taxa, apparently a normal feature of floral development. Spathiphyllum solomonense Nicolson is described in an appendix.     

STRUCTURAL CHANGES IN POPULUS DELTOIDES TERMINAL BUDS AND IN THE VASCULAR TRANSITION ZONE OF THE STEMS DURING DORMANCY INDUCTION

Morphological and anatomical changes in shoots of vigorously growing cottonwood plants (Populus deltoides Bartr.) were studied during dormancy induction in 8-hr short days (SD) and in control plants grown in 18-hr long days (LD). Pronounced structural changes occurred in terminal buds after 4 wk and full dormancy was achieved in 7 wk of SD. Leaf expansion ceased after 5 wk of SD as foliage leaves matured to the terminal bud base at leaf plastochron index 0 (LPI 0). Within the bud, total leaf length (lamina + petiole) decreased and stipule length increased progressively each week; thus, the ratio total leaf length/stipule length decreased rapidly, especially at the position of incipient bud-scale leaves LPI - 1 and LPI - 2. These bud-scale leaves were fully developed by wk 6 and were derived from enlarged stipules and aborted laminae. The full complement of primordia within the bud at the start of SD eventually matured as foliage leaves and the first bud-scale leaf (LPI - 1) was initiated immediately following transfer to SD. Acropetal advance of the primary-secondary vascular transition zone (TZ) was associated with leaf maturation. However, it did not advance throughout the entire vascular cylinder as in LD, but only in those leaf traces serving mature leaves beneath the terminal bud. In both LD and SD treatments the same linear relationship was maintained between LPI of the TZ and LPI of the most recently matured leaf; both parameters simultaneously increased in LD and decreased in SD. Thus, the relationship between leaf maturation and advance of the TZ was maintained irrespective of environment.     

SYSTEMATIC OCCURRENCE OF A SCLEROTIC HYPODERMIS IN ROOTS OF ARACEAE

A survey of the systematic occurrence of a sclerotic hypodermis was made in 91 genera and over 280 species of Araceae. The sclerotic hypodermis in roots consists of a cylinder inside the exodermis, one to five or more cells wide, with thickened, lignified cell walls. The cells are elongated, pitted, and often septate. A sclerotic hypodermis occurs in roots of only eight genera of Araceae, including Culcasia, Montrichardia, Cercestis, Rhektophyllum, Furtadoa, Homalomena, Philodendron, and Anubias. In Philodendron section Philodendron a sclerotic hypodermis is present in aerial and absent from subterranean roots. These results are consistent with other anatomical studies in supporting a transfer of Culcasia from Pothoideae, and Cercestis, Rhektophyllum, and Montrichardia from Lasioideae into an expanded Philodendroideae, as recently proposed by Grayum. Further, these and other anatomical results both provide support for a closer taxonomic association of Culcasia, Cercestis, Rhektophyllum, Philodendron, Montrichardia, Homalomena, Anubias, and Furtadoa than has previously been recognized.     

THE ORGANIZATION OF THE VASCULAR SYSTEM IN THE STEMS OF THE NYMPHAEACEAE. I. NYMPHAEA SUBGENERA CASTALIA AND HYDROCALLIS

The vascular system in the stems of Nymphaea odorata and N. mexicana subgenus Castalia, and N. blanda subgenus Hydrocallis consists of continuing axial stem bundles with eight being the usual number. The stem bundles are concentric and xylem maturation is mesarch. Xylem elements consist of tracheids with spirally or weakly reticulated secondary wall thickenings. The phloem is made up of companion cells and short sieve tube members with simple sieve plates that are nearly transverse. At the node each leaf is supplied with two lateral leaf traces and a median leaf trace. A root trace is also present and supplies a series of adventitious roots borne on the leaf base. Flowers and vegetative buds develop directly from the apical meristem and occupy leaf sites in a single genetic spiral. Each flower or vegetative bud is related to a leaf through specific spatial and vascular association. The related leaf is separated from the related flower by three members of the genetic spiral and occupies an adjacent orthostichy. Vascular tissue for the related flower arises from the inner surfaces of the four stem bundles supplying leaf traces to the related leaf and extends through the pith to the flower or vegetative bud via a peduncle fusion bundle. The vascular system organization in the investigated species of Castalia and Hydrocallis is not typically monocotyledonous or dicotyledonous, nor can it be considered transitional between them. The ontogeny of the vascular system is similar to typical dicotyledons and the investigated species of Nymphaea can, therefore, be considered to represent highly specialized and modified dicotyledons.     

METAMORPHOSIS IN THE ARACEAE

In the Araceae, as in many other monocotyledons, the stem undergoes gradual thickening as successive internodes are produced. Along with this primary thickening of the stem, successive foliage leaves will be larger and often more complex. In many species, in addition to these and other gradual changes, there are abrupt changes, metamorphoses, that occur under certain conditions. Some metamorphoses are the result of endogenous cycling (e.g., Syngonium, Rhektophyllum, Philodendron linnaei), some are a response to changes in conditions in the environment, usually the gain or loss of contact with a substrate (e.g., Syngonium, Monstera, Rhodospatha, Philodendron section Pteromischum), and some occur when the plant reaches a certain level of maturity (e.g., Monstera). Some of these latter changes are associated with a transition from monopodial to sympodial growth (e.g., Philodendron, Anthurium). Metamorphosis enhances the developmental plasticity of the Araceae in two distinct ways: it allows for a more complex relationship between size and shape in the development to maturity, and it allows shoots to engage in dispersal activities and developmental holding patterns when conditions are not suitable for development to the adult form and reproduction.     

LEAF TYPES IN THE ARACEAE

Leaf types in the Araceae are described and classified on the basis of their morphology and functional role. Four classes are recognized on the basis of their association with the initiation of new shoot axes, the continuation of axes, the resting of axes, or the termination and renewal of axes. The basic types are described with the terms leaf, prophyll, mesophyll, bracteole, mesobracteole, cataphyll, and blastophyll. These terms are modified with the terms monopodial, sympodial, proleptic, sylleptic, resting, flagellar, stolon, reduced, and foliage. This represents an unconventional terminology because some of the modifiers refer to the structure of the stem to which the leaves are attached, rather than to the form of the leaf itself. The intent is to draw attention to the impact of shoot organization on leaf form, and to develop a leaf terminology that will aid in describing shoot organization.     

DIVERSITY OF SHOOT MORPHOLOGY IN TYPHONIUM (ARACEAE)

The details of shoot organization, including number of leaves per shoot, position of foliage leaves and cataphylls, position of the lateral continuation shoot, nature of axillary buds and phyllotaxis, and the pattern of shoot extension were observed and compared in five species of Typhonium, Dracunculus vulgaris, Sauromatum venosum, Arum italicum, and Helicodiceros muscivorus, resulting in the recognition of four types of stems. Three of the four types were found in the genus Typhonium. One type was found in Typhonium giganteum and Sauromatum venosum and is presumably the same in Biarum. Another type, found in T. trilobatum, T. blumei, and T. flagelliforme, is distinct in the position of the lateral continuation shoot, which arises from the axil of the n-1 leaf and is adnate to the axis to above the n leaf. Based on the results, two groups, one of which is further subdivided into two subgroups, are recognized in subtribe Arinae (subfam. Aroideae tribe Areae).     

EVOLUTIONARY AND ECOLOGICAL SIGNIFICANCE OF STARCH STORAGE IN POLLEN OF THE ARACEAE

Starch content was qualitatively assessed for pollen of 79 of the 111 currently recognized genera of the family Araceae—one of three monocot families known to exhibit both starchy and starchless pollen. Although 73% of the genera investigated had exclusively starchy pollen, character correlation suggests that starchless pollen is the primitive type for the family Araceae, as well as for monocots in general. Pollen starch content is a highly conservative character at the generic level in Araceae; only a single genus (Schismatoglottis) clearly exhibits both character states. The distribution of starchy pollen among aroid genera is consistent with what have here been termed Bakers' Starch Laws. Aroid pollen below a certain critical diameter—17-25 μm—is almost invariably starchless. Larger pollen is nearly always starchy, except where insect pollinators may use pollen nutritionally. There is strong evidence that the trend from starchless to starchy pollen in Araceae is reversible, according to the constraints imposed by the aforementioned factors.