Trophotagma Enrichment Axes in Poaceae

This review is intended: (1) to interpret and characterize morphological variations observed in the structure of the enrichment axes, located below the terminal inflorescence in Poaceae; and (2) to study the relationship between the intensity of development of such axes and the size of terminal inflorescence. An important reduction in the development of the terminal inflorescence is generally accompanied by a significant development of enrichment axes. It is necessary to adequately characterize these enrichment axes, differentiating them from the terminal inflorescence. Since the intensive development of enrichment axes in synflorescences of many grass genera has caused misinterpretations of the inflorescence structure, to include them as parts of the terminal inflorescence.     

Synflorescences of species related to Schizachyrium condensatum (Poaceae)

The synflorescences of species related to Schizachyrium condensatum are typologically characterized. This species presents floriferous shoots with complex systems of ramification. In all species the inflorescence is polytelic and truncate. The typological pattern has been described and presents differences in the following parameters: length and shape of the synflorescence, paracladia of the trophotagma subzone and short paracladia subzone. A comparative analysis of the variations observed in the structure of the synflorescence is included.     

Morphological architecture of Actinocephalus (Koern.) Sano (Eriocaulaceae-Poales)

Actinocephalus exhibits perhaps more diversity in habit than any other genus of Eriocaulaceae. This variation is largely a result of differences in the arrangement of the paraclades. Based on the analysis of stem architecture of all 25 species of Actinocephalus, the following patterns were established: (1) leaf rosette, with no elongated axis, instead the axillary paraclades originating directly from the short aerial stem, (2) rosette axis continuing into an elongated axis with spirally arranged paraclades, (3) an elongated axis originating from a rhizome, with ramified paraclades, and (4) an elongated axis originating from a short aerial stem, with paraclades arranged in a subwhorl. The elongated axis exhibits indeterminate growth only in pattern 4. Patterns 3 and 4 are found exclusively in Actinocephalus; pattern 1 occurs in many other genera of Eriocaulaceae, while pattern 2 is also found in Syngonanthus and Paepalanthus. Anatomically, each stem structure (i.e., paraclade, elongated axis, short aerial stem, rhizome) is thickened in a distinctive way and this can be used to distinguish them. Specifically, elongated axes and paraclades lack thickening, thickening of short aerial stems results from the primary thickening meristem and/or the secondary thickening meristem. Thickening of rhizomes results from the activity of the primary thickening meristem.     

宽叶兔儿风花序侧生分枝的变异式样以及分枝兔儿风的分类地位问题

宽叶兔儿风Ainsliaealatifolia花序主轴上的侧生分枝通常被描述为由单个或2–4个头状花序聚集而组成的簇生花序。作者在野外和标本馆中仔细研究了该种居群内侧生分枝花序形态的变异,依据其简化和集中的程度在居群内区分出3种花序类型:简单的复合聚伞花序、复杂的复合聚伞花序以及两者之间的过渡类型,具有后面两种类型的个体在居群内少见。这一发现不仅为探讨兔儿风属内各种花序的演化提供了新的线索,而且为理清宽叶兔儿风这一地理广布种的分类学问题提供了新的视野。居群内有些侧生分枝为复杂的复合聚伞花序类型的个体整个花序的主轴折断了,这是由于整个主轴顶端部分被昆虫蛀食或者其他外力造成的。值得注意的是分枝兔儿风Ainsliaealatifoliavar.ramifera的主模式就属于这种情况。分枝兔儿风和宽叶兔儿风原变种之间在分布区、生境以及物候上并没有明显的分化,因此分枝兔儿风被处理成宽叶兔儿风的异名。     

Processes responsible of the structural diversity of the Cyperaceae synflorescence: Hypothetical evolutionary trends

The synflorescence structure of Cyperaceae is revised. It is postulated that the wide diversity shown by the synflorescences can be explained by the analysis of some processes that operate in different ways combined or independently. Twenty-five of such processes are identified and described as responsible for the great diversity detected in the synflorescences of Cyperaceae. Based on these processes, it is possible to recognize the origin of the different types of synflorescences. The paniculodium appears to be the basal unit of inflorescence within the Cyperaceae. Homogenization and truncation possibly are controlling the passage from Juncaceae to Cyperaceae. The diverse processes and the resulting structures were mapped in the phylogeny of the family, demonstrating that the trends of diversification of the synflorescence between the different tribes and within the tribes are mainly reductive.     

Comparative morphology of acrogenous branch systems and phylogenetic considerations. II. Angiosperms

A concept for a primitive angiospermous branch system is given in order to have a starting point for the derivation of the diverse and highly differentiated branch systems observed in contemporary angiosperms. Hitherto Troll's (1964, 1969) comparative study of the synflorescences in this plant group — developed out predominantly on herbaceous plants — was the most comprehensive and sophisticated treatment dealing with branch systems. Unfortunately, the work on tropical tree architecture by Hallé et al. (1978) has no reference to the classical studies of Troll and his pupils. Thus Müller-Doblies and Weberling (1984) emphasized the high degree of terminological incompatibility between the two works. Angiosperms are seen as a monophyletic plant division. Consequently, the branch system of the first primitive angiosperms must be the starting point in the evolution of the abundant diversity of branch systems and growth forms of modern angiosperms. If it is accepted that primitive angiospermous shoots were terminated by a large flower, one may assume, that the reproductive end of the shoot was enriched by paracladia early in evolution, thereby developing a terminal inflorescence instead of a single flower. Thus the primitive shoot unit was divided into a basal vegetative region — the trophotagma, branching retardively — and the reproductive terminal region, the anthotagma, branching simultaneously. It is demonstrated through a selection of different examples, that the construction of such a system possesses the options for several modifications, enabling the evolution of the abundant diversity of branch systems which characterizes contemporary angiosperms.     

Specification of chimeric flowering shoots in wild-type Arabidopsis

Within wild-type Arabidopsis populations, a subset of the plants were found to have a single chimeric shoot on their primary shoot axes. The chimeric shoots were located below the lowest primary-axis flower; and they exhibited features of both flowers and paraclades (lateral flowering shoots). Morphological analyses of chimeric shoots indicated that they developed from single primordia. In each chimeric shoot, the side furthest from the apical meristem was specified as 'flower'—while the side closest to the meristem was specified as 'paraclade'—suggesting that a stimulus from outside the apical meristem can directly induce primordia to develop as flowers. It is concluded that the development of the teratological chimeric shoots resulted from the overlap of the vegetative and floral specification processes within single primordia.     

An assessment of morphogenetic fluctuation during reproductive phase change in Arabidopsis

Background and Aims

Reproductive phase change in Arabidopsis thaliana is characterized by two transitions in phytomer identity, the differentiation of the first elongate internode (bolting transition) and of the first flower (floral transition). An evaluation of the dynamics of these transitions was sought by examining the precision of the corresponding phytomer identity changes.

Methods

The length of the first elongate internode and the frequency of chimeric inflorescence structures, e.g. paraclades not subtended by a leaf (no-leaf/paraclades) and flowers subtended by a bract (bract/flowers), were measured in the Wassilewskija (Ws) accession and 47 early flowering mutants under a wide range of photoperiods. The impact of photoperiodic perturbations applied to Ws plants at different times of development was also evaluated.

Key Results

In Ws, both types of characters were remarkably constant across photoperiods in spite of a high degree of interindividual variability. Bract/flowers were not normally produced in Ws, but they were observed in conditions that suggest enhanced light signalling, e.g. in response to continuous light perturbations and in mutants with reduced hypocotyl elongation. In contrast, no-leaf/paraclades were normally present in approx. 20 % of Ws plants, and their frequency was increased in conditions that suggest reduced light signalling, e.g. in mutants with altered specification of long-day responses. The length of the first elongate internode was unrelated to the rate of stem elongation and to the regulation of reproductive phase change.

Conclusions

Bract/flowers and no-leaf/paraclades corresponded to opposite effects on the floral transition that reflected different dynamics of progression to flowering. In contrast, the length of the first elongate internode was only indirectly related to the regulation of reproductive phase change and was mainly dependent on global morphogenetic constraints. This paper proposes that morphogenetic variability could be used to identify critical phases of development and characterize the canalization of developmental patterns.     

Resistance of insecticide-treated foam board insulation against the eastern subterranean termite and the Formosan subterranean termite (Isoptera: Rhinotermitidae)

Three foam board types, one untreated control, one containing 2,000 ppm disodium octaborate tetrahydrate (DOT), and one containing 1,000 ppm deltamethrin, were exposed to field populations of the eastern subterranean termite, Reticulitermes flavipes (Kollar), and the Formosan subterranean termite, Coptotermes formosanus Shiraki. There was no significant difference in termite damage between foam boards treated with 2,000 ppm DOT and the untreated control. Form boards containing 1,000 ppm deltamethrin were not damaged by R. flavipes, whereas only minor damage occurred after exposure to C. formosanus.     

Inflorescence structure in Rhynchospora Vahl (Cyperaceae)

We investigated the inflorescence structure of Rhynchospora following the methodology and terminology of Troll's school, with the objective of providing a characterization of the inflorescence suitable to evaluate the processes responsible for the diversity observed. Homogenization of inflorescence structures may occur fully or partially. In the first case, all branches of the inflorescence are homogeneous, while in the later, distal and proximal parts of the inflorescence bear homogeneous branches, but in the middle portions of the inflorescence non-homogeneous branches exist. Other characters leading to different forms of inflorescences are branching degree, internode elongation along the main axis of the synflorescence, degree of epipodium elongation of distal paraclades, development of bract and prophyll, and development of prophyllar paraclades. We identified three main types of inflorescences: (1) partially homogeneous paniculodia, (2) partially homogeneous capitate heads, and (3) a fully homogeneous capitate head. Within the first type, four subtypes were also recognized. Finally, we discuss how these processes can operate to produce the variation of the inflorescence shape.     

Heat tolerance and acclimation capacity in subterranean arthropods living under common and stable thermal conditions

Cave‐dwelling ectotherms, which have evolved for millions of years under stable thermal conditions, could be expected to have adjusted their physiological limits to the narrow range of temperatures they experience and to be highly vulnerable to global warming. However, most of the few existing studies on thermal tolerance in subterranean invertebrates highlight that despite the fact that they show lower heat tolerance than most surface‐dwelling species, their upper thermal limits are generally not adjusted to ambient temperature. The question remains to what extent this pattern is common across subterranean invertebrates. We studied basal heat tolerance and its plasticity in four species of distant arthropod groups (Coleoptera, Diplopoda, and Collembola) with different evolutionary histories but under similar selection pressures, as they have been exposed to the same constant environmental conditions for a long time. Adults were exposed at different temperatures for 1 week to determine upper lethal temperatures. Then, individuals from previous sublethal treatments were transferred to a higher temperature to determine acclimation capacity. Upper lethal temperatures of three of the studied species were similar to those reported for other subterranean species (between 20 and 25°C) and widely exceeded the cave temperature (13–14°C). The diplopod species showed the highest long‐term heat tolerance detected so far for a troglobiont (i.e., obligate subterranean) species (median lethal temperature after 7 days exposure: 28°C) and a positive acclimation response. Our results agree with previous studies showing that heat tolerance in subterranean species is not determined by environmental conditions. Thus, subterranean species, even those living under similar climatic conditions, might be differently affected by global warming.     

<Emphasis Type="Italic">Miconia rondoniensis</Emphasis> (Melastomataceae), a new species from the Southern Amazon of Brazil

Miconia rondoniensis occurs in secondary upland (terra firme) moist open forest in the Brazilian states of Rondônia, Mato Grosso and Amazonas. It can be recognized by the tree habit, large leaves with rounded to obtuse or subcordate bases and 5+2 basal veins (five main veins plus a faint submarginal pair), large inflorescences with proximal paraclades bearing quaternary branches and distal paraclades bearing tertiary branches, persistent hemi-elliptic bracteoles, short persistent calyx lobes, and capitate stigmas. The most striking feature of this new species is the thick pubescence on the branches and petioles, composed of trichomes having a long axis and short arms at the base, but inflated and lacking arms at the apex, resulting in a sorus-like surface covered with globose structures.     

Genome‐wide adaptive evolution to underground stresses in subterranean mammals: Hypoxia adaption,immunity promotion,and sensory specialization

Life underground has provided remarkable examples of adaptive evolution in subterranean mammals; however, genome‐wide adaptive evolution to underground stresses still needs further research. There are approximately 250 species of subterranean mammals across three suborders and six families. These species not only inhabit hypoxic and dark burrows but also exhibit evolved adaptation to hypoxia, cancer resistance, and specialized sensory systems, making them an excellent model of evolution. The adaptive evolution of subterranean mammals has attracted great attention and needs further study. In the present study, phylogenetic analysis of 5,853 single‐copy orthologous gene families of five subterranean mammals (Nannospalax galili, Heterocephalus glaber, Fukomys damarensis, Condylura cristata, and Chrysochloris asiatica) showed that they formed fou distinct clusters. This result is consistent with the traditional systematics of these species. Furthermore, comparison of the high‐quality genomes of these five subterranean mammalian species led to the identification of the genomic signatures of adaptive evolution. Our results show that the five subterranean mammalian did not share positively selected genes but had similar functional enrichment categories, including hypoxia tolerance, immunity promotion, and sensory specialization, which adapted to the environment of underground stresses. Moreover, variations in soil hardness, climate, and lifestyles have resulted in different molecular mechanisms of adaptation to the hypoxic environment and different degrees of visual degradation. These results provide insights into the genome‐wide adaptive evolution to underground stresses in subterranean mammals, with special focus on the characteristics of hypoxia adaption, immunity promotion, and sensory specialization response to the life underground.     

Bi-directional inflorescence development inArabidopsis thaliana: Acropetal initiation of flowers and basipetal initiation of paraclades

In this study we investigated Arabidopsis thaliana (L.) Heynh. inflorescence development by characterizing morphological changes at the shoot apex during the transition to flowering. Sixteen-hour photoperiods were used to synchronously induce flowering in vegetative plants grown for 30 d in non-inductive 8-h photoperiods. During the first inductive cycle, the shoot apical meristem ceased producing leaf primordia and began to produce flower primordia. The differentiation of paraclades (axillary flowering shoots), however, did not occur until after the initiation of multiple flower primordia from the shoot apical meristem. Paraclades were produced by the basipetal activation of buds from the axils of leaf primordia which had been initiated prior to photoperiodic induction. Concurrent with the activation of paraclades was the partial suppression of paraclade-associated leaf primordia, which became bract leaves. The suppression of bract-leaf primordia and the abrupt initiation of flower primordia during the first inductive photoperiod is indicative of a single phase change during the transition to flowering in photoperiodically induced Arabidopsis. Morphogenetic changes characteristic of the transition to flowering in plants grown continuously in 16-h photoperiods were qualitatively equivalent to the changes observed in plants which were photoperiodically induced after 30 d. These results suggest that Arabidopsis has only two phases of development, a vegetative phase and a reproductive phase; and that the production of flower primordia, the differentiation of paraclades from the axils of pre-existing leaf primordia and the elongation of internodes all occur during the reproductive phase.     

STUDIES ON THE ECOLOGICAL AND PHYSIOLOGICAL SIGNIFICANCE OF AMPHICARPY IN GYMNARRHENA MICRANTHA (COMPOSITAE)

The 2 types of fruit (aerial and subterranean) borne by the dwarf desert annual Gymnarrhena micrantha were compared with regard to their responses to factors affecting their formation, dispersal, germination and seedling mortality. The 2 types of fruit differed markedly in several respects. In comparison with the subterranean fruits, the aerial ones are much smaller and more numerous, but the formation of the inflorescence in which they develop is more dependent on a favorable supply of soil moisture. The aerial fruits are dispersed by wind, after becoming detached by a complex series of hygroscopic movements which involve several organs and tissues, while the subterranean fruits never leave the dead parent plant, germinating right through its tissues. Germination of the subterranean fruits starts after a shorter incubation period and is less temperature-dependent in both light and dark. Light stimulated germination of both types of fruit, increasing their germination rates and final percentages, but not affecting the duration of the incubation period. In the subterranean fruits, the rate of germination was equally stimulated by light over the entire temperature range, with a well-defined optimum at 15 C in both light and dark. In the aerial fruits, the same optimum was found only in the light, rates in darkness increasing with decreasing temperatures. In the aerial fruits, alternations of light and dark were more favorable to germination than either continuous light or dark, the full effect being obtained with a single 8-hr or 16-hr light period, provided it was preceded by 16 or 8 hr of darkness, respectively. Similar reactions to combinations of light and dark were not observed in the subterranean fruits. Seedlings developing from the subterranean fruits were much larger, but grew at a relatively much slower rate than those from aerial fruits. The former were distinctly more tolerant of unfavorable soil-moisture regimes, such as low moisture supply and drought. It was concluded that the 2 types of fruit serve 2 distinct functions in the biology of the plant. The aerial fruits are adapted to the function of increasing the distribution of the species within suitable habitats, while the subterranean fruits are adapted to increasing the probability of the survival of the species.     

Life History Evolution in Amphicarpic Plants

Abstract Plants with dimorphic flowers or seeds provide excellent material for the study of life history evolution because the dimorphism often involves measurable differences in morphology, size, number, or genetic relatedness. For amphicarpic plants, the proportion of aerial: subterranean morphs produced is highly variable (from 0 to > 100) and related to both environmental and genetic factors. Plants from aerial seeds produce lower ratios of aerial: subterranean morphs than those from subterranean seeds. Despite substantial variation, subterranean seeds are generally heavier than aerial seeds (but fewer) and produce vigorous seedlings with high survivorship and high fitness. Adaptive advantages of subterranean seeds include retention of offspring in favorable parental microhabitats, protection of seeds from herbivory, predation, or fire, and avoidance of desiccating conditions on the soil surface; potential disadvantages include lack of gene exchange, high energy costs, limited dispersal, and sibling competition. For the few species studied, aerial reproduction is more plastic than subterranean reproduction and more likely to be affected by environmental conditions. Quantitative genetic analyses of a population of the annual grass Amphicarpum purshii have revealed lower heritabilites for subterranean relative to aerial reproductive traits. Subterranean seed number and mass show genetic correlations with shoot mass while aerial seed number and mass do not; seed set percentages of both seed types as well as percentage allocation to both reproductive morphs show negative genetic correlations with shoot mass. In this Amphicarpum population, directional selection on shoot mass may indirectly select for increased subterranean (but not aerial) seed output.     

The Effect of the Apical Bud on the Growth of Lateral Buds on Subterranean Shoots

The influence of the apical bud on the growth of the lateral buds on subterranean shoots was studied in Stachys sieboldiiMig. and Helianthus rigidus(Gass.) Desv. Removing and damaging the apical parts of subterranean shoots or their treatment with 2% chlorocholine chloride shoot enhanced shoot branching. The response to light of the apical bud was invariably negative: the stolons, which came out or were extracted from the soil, grew back into the ground (negative phototropism). The response to light of lateral buds was autonomous and depended on the conditions of their initiation. The lateral buds developed in darkness manifested negative phototropism when withdrawn from the soil and exposed to the light, whereas the buds developed in the light showed positive phototropism. The author concludes that the concept of apical dominance, thoroughly studied in aboveground shoots, is also valid for subterranean shoots. However, in contrast to the former, in the latter case, the apical bud does not control the growth orientation of the lateral buds.     

Consequences of a mixed reproductive system in the hog peanut,Amphicarpaea bracteata, (Fabaceae)

Summary The forest annual, Amphicarpaea bracteata L. can reproduce via aerial chasmogamous, aerial cleistogamous, and subterranean cleistogamous flowers. Both plant size and light intensity influenced the utilization of the three modes of reproduction. chasmogamous and aerial cleistogamous flower number and the ratio of chasmogamous flowers to the total number of aerial flowers increased with plant size. The latter demonstrated a shift to xenogamy and outbreeding in larger plants. Light intensity indirectly influenced reproductive modes through its infuence on plant size. Seed set by both types of aerial flowers was low and unrelated to plant size. Subterranean seed number and the total dry weight of subterranean seeds per plant increased with size. The subterranean seeds of Amphicarpaea bracteata are thirty-four times larger than the aerial seeds (fresh weight). Under field conditions, subterranean seeds had greater germination after one year than acrial seeds. The plants arising from subterranean seeds were significantly larger and more fecund than those from aerial seeds. Seeds produced by aerial cleistogamous, hand selfpollinated chasmogamous, and naturally pollinated chasmogamous flowers had equivalent germination rates and produced plants of equal size and fecundity. This suggests that the outbred progeny from chasmogamous flowers have no advantage over the inbred progeny from aerial cleistogamous flowers.     

Cell division patterns in the apices of subterranean axis and aerial shoot of Psilotum nudum (Psilotaceae): morphological and phylogenetic implications for the subterranean axis

The cell division pattern in the apical meristem of Psilotum nudum was examined using epi-illumination microscopy and a paraffin method. In the subterranean axis, about half of the derivative cells of the apical cell produce tetrahedral daughter apical cells by the first three or more oblique divisions. Roughly half of these apical cells give rise to the apical meristems of axes, whereas the other half do not. Various relative activities of the mother and daughter apical cells give rise to disordered branching patterns. In the ill-organized apical meristem as well as the leafless and capless structure, the Psilotum subterranean axis differs from the basic organs of vascular plants such as stem and root and seems to be an independent organ. The cell division pattern characteristic of the subterranean axis persists in the young unbranched aerial shoots, although fewer daughter apical cells are produced. Dichotomous branching of the aerial shoots, as in a variety of organs of pteridophytes, involves loss of the mother apical cell followed by appearance of two daughter apical cells.     

RESOURCE ALLOCATION AND REPRODUCTION IN POPULATIONS OF AMPHICARPUM PURSHII (GRAMINEAE)

Five New Jersey populations of Amphicarpum purshii (an annual panicoid grass) were investigated as to total allocation of biomass to reproduction and its distribution between aerial chasmogamous and subterranean cleistogamous inflorescences. Germination of both types of seed was documented, for the first time, in not only the field but under three laboratory temperature regimes. Overall, approximately 29 % of the shoot biomass was allocated to reproduction. On a population basis, subterranean inflorescences accounted for 37–100 % of the reproductive biomass, and these seed (5 × heavier but fewer in number) were the source of most surviving seedlings. Subterranean spikelets contained caryopses significantly more frequently than aerial spikelets. The ratio of the number of viable aerial seed to the number of viable subterranean seed increased from 0/4 to 4/2 with recentness and/or frequency of disturbance. As a pioneer species in secondary succession, Amphicarpum purshii produces a larger subterranean propagule, with greater seedling vigor and a higher probability of local reproductive success, and also a smaller aerial propagule in larger numbers with potentially greater genetic variability.