三倍体丹参杂交种的花粉形态研究

三倍体丹参是以二倍体丹参为父本、人工染色体加倍的四倍体白花丹参为母本杂交选育的杂交种。为深入了解三倍体丹参花粉的特性,以及为三倍体种质利用提供孢粉学依据,该文以二倍体丹参为对照,研究了三倍体丹参杂交种花粉的形态变异规律。利用光学显微镜和扫描电镜对二倍体和三倍体丹参的花粉萌发沟、外壁纹饰、花粉粒形状等特征进行了显微和超微形态观察,综合进行了花粉形态差异比较,并对花粉大小和形状数据进行了差异显著性分析和正态检验。结果表明:(1)二倍体丹参为6沟花粉,三倍体花粉萌发沟有6沟和8沟两种类型,沟内疣状颗粒分布不匀,出现畸形萌发沟。(2)二倍体和三倍体花粉外壁均为网状雕纹。二倍体花粉网眼内具多个多边形穿孔,穿孔大; 6沟和8沟两种类型的三倍体花粉网眼无穿孔或仅有几个小穿孔,6沟和8沟花粉的外壁雕纹相同。(3)三倍体花粉的极轴长(P)和赤道宽(E)均值显著小于二倍体花粉,花粉大小呈偏正态分布,P*E的差异系数大于二倍体花粉,且有极值存在。三倍体和二倍体丹参的萌发沟和雕纹存在差异,而花粉形状差异不显著。综上结果表明三倍体丹参花粉在倍性效应和杂合性的双重影响下发生了形态变异,且有多种形态变化。     

Pollen anomaly - a clue to natural hybridity in Argemone (Papaveraceae)

Studies in the pollen morphology of Argemone mexicana (a diploid species, 2n=28), A. ochroleuca (a tetraploid, 2n=56), the wild populations of which growing side by side, and of a triploid (2n=42) observed in the population of A. ochroleuca, have been carried out using light and scanning electron microscope. Pollen grains in both the species are 3 (-4)-zonocolpate with reticulate exine surface, and are distinguishable only on the basis of pollen size, the tetraploid species having larger pollen grains. In the triploid plant, a large number of grains deviates from the normal 3-zonocolpate condition showing a number of apertural types, such as 1-aperturate-operculate, 2-colpate, 3-syncolpate, 4-zonocolpate and spiraperturate. Besides, planaperturate 3-colpate-trilobed grains also occur. The triploid is also characterised by 'minigrains' (size 13-19 μm), a feature not observed in A. mexicana and A. ochroleuca. Since these species are reported to be genetically close, the occurrence of the triploids (forming nearly 5% of the population of A. ochroleuca) indicates the possibility of a natural hybridization between A. ochroleuca and A. mexicana, which is well pronounced on its anomalous pollen features serving as marker for such a hybrid occurring in nature.     

Pathways to polyploidy: indications of a female triploid bridge in the alpine species <Emphasis Type="Italic">Ranunculus kuepferi</Emphasis> (Ranunculaceae)

Polyploidy is one of the most important evolutionary processes in plants. In natural populations, polyploids usually emerge from unreduced gametes which either fuse with reduced ones, resulting in triploid offspring (triploid bridge), or with other unreduced gametes, resulting in tetraploid embryos. The frequencies of these two pathways, and male versus female gamete contributions, however, are largely unexplored. Ranunculus kuepferi occurs with diploid, triploid and autotetraploid cytotypes in the Alps, whereby diploids are mostly sexual, while tetraploids are facultative apomicts. To test for the occurrence of polyploidization events by triploid bridge, we investigated 551 plants of natural populations via flow cytometric seed screening. We assessed ploidy shifts in the embryo to reconstruct female versus male gamete contributions to polyploid embryo and/or endosperm formation. Seed formation via unreduced egg cells (B_III hybrids) occurred in all three cytotypes, while only in one case both gametes were unreduced. Polyploids further formed seeds with reduced, unfertilized egg cells (polyhaploids and aneuploids). Pollen was highly variable in diameter, but only pollen >27 μm was viable, whereby diploids produced higher proportions of well-developed pollen. Pollen size was not informative for the formation of unreduced pollen. These results suggest that a female triploid bridge via unreduced egg cells is the major pathway toward polyploidization in R. kuepferi, maybe as a consequence of constraints of endosperm development. Triploids resulting from unreduced male gametes were not observed, which explains the lack of obligate sexual tetraploid individuals and populations. Unreduced egg cell formation in diploids represents the first step toward apomixis.     

Characteristics of pollen from natural triploid Betula hybrids

Birch has a key role in the Holocene vegetation history of northern Europe and in sub‐arctic climates dwarf birch and tree birch co‐exist. In Iceland, triploid hybrids between diploid Betula nana (dwarf birch) and tetraploid B. pubescens (downy birch) are common and therefore likely to contribute to pollen deposition. Pollen from 22 triploid trees/shrubs from ten woodlands in Iceland was examined and its size and shape compared with pollen from the parental species. The mean diameter of pollen grains from the triploid hybrids was not statistically different from that of B. nana pollen, but was significantly smaller than the mean value of B. pubescens pollen. On the contrary, the size of the vestibulum was similar to that of B. pubescens, which was significantly greater than that of B. nana, and therefore the diameter‐pore depth ratio was lower than the values from either species. The pattern of size distribution within plants indicated that triploid hybrids might have produced two sizes of triporate pollen grains, but the small B. nana size was far more prevalent than the larger B. pubescens size. Several anomalies in pollen morphology were common among the hybrid pollen grains: four or more pores were the most frequent type of abnormality. Characteristics of the pollen of triploid Betula hybrids, especially structural anomalies, may provide a means to reveal periods of interspecific hybridisation in the analysis of sub‐fossil pollen.     

Ploidy distribution and DNA content variations of <Emphasis Type="Italic">Lonicera caerulea</Emphasis> (Caprifoliaceae) in Japan

Ploidy level and geographical distribution were investigated in Japanese Lonicera caerulea L. Flow cytometric analysis revealed the presence of DNA diploid and DNA tetraploid plants in Japan. Chromosome observation confirmed that diploid and tetraploid plants showed 2n = 2x = 18 and 2n = 4x = 36, respectively. The DNA diploid populations were found only in lowland mires, Betsukai, Bekanbeushi, Kushiro and Kiritappu located in eastern Hokkaido. On the other hand, DNA tetraploid populations were distributed in a wide area of Hokkaido, and mainland of Japan. The habitats of DNA tetraploid plants were lowland to alpine region. The DNA content measurement with flow cytometry revealed significant differences in the relative DNA contents among DNA tetraploid populations. The relative DNA content within DNA tetraploid populations varied 1.157-fold at maximum, and might correlate with altitude indicating that DNA contents were smaller as altitude increases. The wide area of distribution in various environments of DNA tetraploid plants suggested the adaptability of the tetraploid plants. Although diploid and tetraploid populations were found, no triploid was detected, indicating crossing difficulty between diploid and tetraploid as confirmed by crossing experiment.     

Production of tetraploid plants of non apomictic citrus genotypes

Ploidy manipulation in Citrus is a major issue of current breeding programs aiming to develop triploid seedless mandarins to address consumer demands for seedless fruits. The most effective method to obtain triploid hybrids is to pollinate tetraploid non apomictic cultivars with pollen of diploid varieties. Such non apomictic tetraploid lines are not found in the citrus germplasm and need to be created. In this work we describe a new methodology based on in vitro shoot-tip grafting combined with treatment of the micro-grafted shoot-tip with colchicine and oryzalin to achieve chromosome doubling and a dechimerization procedure assisted by flow cytometry. Stable tetraploid plants of Clemenules, Fina and Marisol clementines and Moncada mandarin have been obtained directly from shoot tip grafting combined with colchicine and oryzalin treatments or after dechimerization of mixoploids plants (2x–4x). These stable tetraploid plants have been used in 4x × 2x hybridizations, to recover over 3,250 triploid hybrids in 3 years.     

Natural hybridisation in birch: triploid hybrids between Betula nana and B. pubescens

Hybridisation between diploid (2n=28) dwarf birch Betula nana L. and tetraploid (2n=56) downy birch B. pubescens Ehrh. has occurred in natural populations in Iceland. About 10% of birch plants randomly collected are triploid (2n=42) hybrids. Ribosomal gene mapping on chromosomes and genomic in situ hybridisation confirms the hybridity. However, the triploid hybrids are not morphologically distinct, i.e. they are not different from diploid and tetraploid birch plants that have intermediate morphology. The triploid hybrids have evidently played an important role in driving bi-directional gene flow between these two species. This paper reviews the extent of interspecific hybridisation in selected birch woodland populations and discusses the significance of natural hybridisation and introgression in birch.     

Ecological factors influencing tetraploid establishment in snow buttercups (Ranunculus adoneus , Ranunculaceae): minority cytotype exclusion and barriers to triploid formation

Polyploid speciation is an ongoing, important source of angiosperm diversity. However, the barriers to polyploid speciation and mechanisms of establishment remain poorly understood for all but a few species. Several factors likely to have influenced tetraploid establishment, including barriers to triploid formation, consequences of mixed-ploidy pollen loads, and the reproductive success of the minority cytotype, were examined in snow buttercups (Ranunculus adoneus) through a series of pollination and transplant experiments. Tetraploid snow buttercups do not have stigmatic barriers to pollen from diploid plants, nor does pollen from tetraploid plants have an advantage over pollen from diploids when on tetraploid stigmas. Tetraploid plants transplanted into a diploid population produced 50% fewer seeds than tetraploid plants in a tetraploid population. Intrinsic barriers to triploid formation were relatively weak, but few triploid seeds formed when mixed-ploidy pollen was present. Fecundity of triploid plants was very low, and no tetraploid offspring resulted. These results indicate that in snow buttercups, a triploid plant will contribute 0.8% of the tetraploid seeds of a minority tetraploid plant making it a minor contributor to the demographics of tetraploid establishment. The reproductive costs facing minority cytotype plants may explain the previously observed spatial segregation in snow buttercups.     

CHROMOSOME ANALYSIS OF HYBRIDS BETWEEN 4N GRINDELIA CAMPORUM AND 2N G. GRANDIFLORA (COMPOSITAE)

Morphology and meiosis are described in four progeny plants resulting from tetraploid Grindelia camporum Greene (2n = 24) from California pollinated by diploid G. grandiflora Hook. (2n = 12) from Coahuila, Mexico. Three of the four progeny were tetraploid, morphologically like the pistillate parent, and had metaphase I chromosome configurations which included quadrivalents and a complementary number of bivalents. They are considered to have resulted from selfing. The fourth plant was triploid (2n = 18) andmorphologically intermediate between the parents. Chromosome configurations in the triploid were variable with univalents, ring and rod bivalents, trivalents and pentavalents. These two species are considered related through an ancestor with a basic genome, but are separated cytologically by polyploidy and by two distinct chromosomal interchanges that explain the configurations observed in the triploid hybrid.     

Production and viability of unreduced gametes in triploid interspecific blueberry hybrids

Summary Three triploid (2n=3x=36) blueberry hybrids were obtained by hand-pollinating approximately 7,000 flowers of tetraploid highbush blueberry cultivars (based on Vaccinium corymbosum L.) with pollen from the diploid species V. elliottii Chapm. Meiotic analysis of these triploids revealed trivalents, bivalents and univalents in all metaphase I cells, with lagging chromosomes evident at anaphase I. Pollen of the three triploids was mostly aborted and did not stain with acetocarmine. However, the three triploids did produce from 0.9%–1.3% giant pollen grains that stained with acetocarmine and were present as monads, dyads or triads, rather than the normal tetrads. Pollination of 10,853 flowers of hexaploid V. ashei Reade cultivars with pollen from the triploids produced 266 berries, which averaged fewer than two fully-developed seeds per berry. One triploid clone showed partial female fertility when crossed to hexaploids, self-pollinated, or intercrossed with other triploids. Ploidy levels of the resulting hybrids were determined.Florida Agricultural Experiment Station Journal Series No. 8672     

The early ontogeny of embryoids and callus from pollen and subsequent organogenesis in anther cultures of Datura metel and rice

Summary Haploidy induction through anther culture has been examined in Datura metel and rice with a view to tracing the precise sequence of development of the pollen, either directly or through an intervening callus, into an embryo and seedling. In D. metel, the vegetative cell of the young pollen grain assumes the major role in formation of embryos whereas the generative cell and its few derivatives degenerate. Embryos and seedlings arising directly from pollen without an intervening callus phase always proved to be haploids, whereas those differentiating from pollen-derived callus gave haploid, diploid and even triploid plants. Cytological analysis of callus tissue showed cells of various ploidy levels ranging from haploid to triploid, and in rare instances even with higher chromosome numbers.In rice anther cultures the embryoids arose from an initial callus phase. Of 15 different rice cultivars tried, only four produced a callus, and in only one, was there differentiation of plants, both haploid and diploid ones. Among other species tried, egg plant has also yielded plantlets through a callus phase whereas only callus production has been achieved in jute, tea and petunia. No response has been obtained in wheat, maize, cotton and coconut.Coconut milk (CM) appears to be the most important component of the medium for the initial induction of embryoids and callus in anther cultures of most of the species tried. However, further growth and differentiation of plants may require a simpler medium; in D. metel, continued culture on CM led to dedifferntiation.Dedicated to the memory of the late Dr. J. P. Nitsch.     

THE PROBLEM OF DRYOPTERIS LEEDSII

Walker , S. (U. Liverpool, England.) The problem of Dryopteris leedsii. Amer. Jour. Bot. 49(9): 971–974. Illus. 1962.—Investigation of 12 plants of ‘Dryopteris leedsii‘ from the type locality in Harford County, Maryland, has revealed 3 cytological entities, diploid, triploid and tetraploid. The diploid and triploid types exhibit hybrid characters and are sterile, while the tetraploid is a fertile species. The relationship between the 3 entities remains a problem, but the possibilities are discussed.     

Induced single fertilization in maize

 Bicellular pollen with one vegetative nucleus and one diploid arrested generative cell (”monospermic” pollen) was induced by trifluralin treatment of diploid maize plants at 7–9 days before flowering. The arrested generative cell (seemingly a diploid sperm cell) fused with the central cell of diploid plants and produced shriveled endosperm resembling that of a 2n×4n cross in maize. Dual pollination experiments with a purple embryo marker revealed single fertilization events in which the union of one sperm cell with the egg occurs but there is no union of a second sperm cell with the central cell. Singly fertilized ovules survived at least 4 days. Furthermore, many viable triploid plants were obtained. This technique therefore appears to have the potential for manipulating ploidy level in crops and may become useful in investigating fertilization mechanisms of angiosperms. Received: 1 October 1996 / Revision accepted: 8 January 1997     

A CYTOGEOGRAPHIC STUDY OF ERIOPHYLLUM LANATUM (COMPOSITAE,HELENIEAE)

Analysis of 368 plants derived from 239 natural populations showed that this taxonomically perplexing and wide-ranging species-complex consists of diploids (n = 8), tetraploids, hexaploids and octoploids. Microsporocytes were the source of most of the chromosome counts. Meiosis was basically regular. Multivalent formation was uncommon, but 11 % of all the plants examined had one or more full-sized extra chromosomes. The frequency of plants with extra chromosomes varied significantly among the taxa, from 0 (five varieties) to over 20 % (two varieties). Except in one instance, where one population yielded a diploid and a triploid, different ploidy levels were not found in the same population. The frequency of diploid, tetraploid, hexaploid and octoploid populations was, respectively, 71, 22, 4 and 2%. Variety obovatum appears to be exclusively diploid, and var. aphanactis exclusively tetraploid. Diploids and one or more polyploid levels occurred in the other taxa. No correlation was found between polyploidy and geological history, soils, topography or climate, nor were the polyploids more widely distributed than the diploids. Some of the polyploid populations seem to have been derived from inter-varietal hybridizations, but others do not. The complex has a “pillar” structure in which 10 diploid taxa support a three-level polyploid superstructure. The available evidence suggests that the major role of polyploidy here has been to stabilize the products of intra- and inter-varietal hybridizations.     

In vitro embryo rescue culture of F<Subscript>1</Subscript> progenies from crosses between diploid and tetraploid grape varieties

The purpose of this study was to understand factors affecting in vitro embryo rescue culture from hybrids between diploid and tetraploid varieties of grape in creation new triploid germplasm resources. The effects of different media, removal ages of immature seeds and reciprocal crosses of parents on the germination and seedling survival of immature seeds from crosses between diploid and tetraploid grape varieties by in vitro embryo rescue culture were investigated. The results indicated that the medium consisting of NN-1969 + IAA 1.75 mg l⁻¹ + GA₃ 0.35 mg l⁻¹ + CH 400 mg l⁻¹ + AC 2.0 g l⁻¹ was better than other media. The optimal removal age of immature seeds for the best development of embryos was 35–45 days after pollination (DAP). The percentage of germination (PG) for immature seeds and the percentage of seedling survival (PSS) for immature seeds for diploid varieties used as female parents were 10.72% and 4.35% higher than when tetraploid varieties were used as female parents respectively. A total of 41 hybrid progenies from eight combinations were obtained, made up of 17 diploid, 9 tetraploid, 14 aneuploid, and 1 triploid progeny as determined by root-tip chromosome identification. The triploid progeny was from Fujiminori (2n = 4x = 76) × Jingxiu (2n = 2x = 38). These results implied that it was feasible to extend the hybridization range of grape and to create new germplasm resources by in vitro embryo rescue based on the conventional hybridization. The NN-1969 medium supplemented with GA₃ and IAA was more propitious to the development of immature seeds sampled at about 45 DAP. It was easier to obtain plants using diploid as female parent, but triploid progeny was only obtained using tetraploid as female parent.     

Use of 2n pollen-producing triploid hybrids to introduce tetraploid Mexican wild species germ plasm to cultivated tetraploid potato gene pool

Summary Tetraploid (2n=4x=48) 2EBN Mexican wild species in the series Longipedicellata, which consists of Solanum fendleri, S. hjertingii, S. papita, S. polytrichon, and S. stoloniferum, were crossed with two 2EBN cultivated diploid (2n=2x=24) clones. The resulting triploid hybrids (2n=3x=36) produced 2n pollen (triplandroids) by the mechanism of parallel orientation of anaphase II spindles. The percentage of stainable pollen in 520 triploids ranged between 0 and 23.5%, with a mean of 2.7%. Triploids producing between 13.0 and 23.5% stainable pollen were crossed as staminate parents to the tetraploid cultivars, resulting in abundant pentaploid (2n=5x=60) and near-pentaploid hybrid progeny. Crosses of triploids with lower percentage of stainable pollen as pollen parent to the tetraploid cultivars did not yield fruit, unless rescue pollen from a tetraploid cultivar was added 2 days later. Pentaploid hybrids were selected among selfed tetraploid progenies using morphological and isoxyme markers transmitted from their cultivated diploid parents. These pentaploid hybrids were vigorous and had uniformly sterile pollen. They were female fertile and were crossed with tetraploid cultivars, yielding an average of 19 seeds per fruit. Triplandroids provide the opportunity of transferring 2EBN tetraploid Mexican wild species in the series Longipedicellata germ plasm into the 4EBN cultivated potatoes.Cooperative investigations of the ARS, USDA, and the Washington State University Agricultural Research Center, Prosser, WA 99350, USA. H/LA Paper No. 90-03, College of Agriculture and Home Economies Research Center, Washington State University, Pullman, WA 99164, USA     

GENETIC CONSEQUENCES OF AUTOPOLYPLOIDY IN TOLMIEA (SAXIFRAGACEAE)

Although there is an extensive literature on the genetic attributes of allopolyploids, very little information is available regarding the genetic consequences of autopolyploidy in natural populations. We therefore addressed the major predicted genetic consequences of autopolyploidy using diploid and tetraploid populations of Tolmiea menziesii. Individual autotetraploid plants frequently maintain three or four alleles at single loci: 39% of the 678 tetraploid plants exhibited three or four alleles for at least one locus. Heterozygosity was also significantly higher in autotetraploid populations than in diploid populations: H_° = 0.070 and 0.237 in diploid and tetraploid Tolmiea, respectively. Most of the genetic diversity in T. menziesii is maintained within populations (ratio of gene diversity within populations to mean total genetic diversity = 0.636). The total genetic diversity due to differentiation between the two cytotypes is only 0.055. Such a low degree of differentiation between cytotypes would be expected between a diploid and its autotetraploid derivative. Most diploid and all tetraploid populations examined are in genetic equilibrium. Diploid and tetraploid Tolmiea share three or four alleles at six of eight polymorphic loci. This suggests that either autotetraploid Tolmiea was formed via crossing of genetically different diploids (perhaps via a triploid intermediate) or autopolyploidy occurred more than once in separate individual plants, followed by later crossing of autotetraploids.     

A CYTOGENETIC ANALYSIS OF CERTAIN POLYPLOIDS IN GRINDELIA (COMPOSITAE)

Two diploid taxa, Grindelia procera and G. camporum, and 3 tetraploid ones, G. camporum, G. hirsutula, and G. stricta, have been studied to ascertain their interrelationships. Meiosis in diploid parental strains was regular, the common chromosome configuration being 5 rod bivalents and 1 ring bivalent. The average chiasmata frequency per chromosome was 0.60. Pollen fertility was about 90% in all strains examined. Diploid interspecific hybrids had normal meiosis with an average chiasmata frequency of 0.56 per chromosome. No heterozygosity for inversions or interchanges was detected, and pollen fertility was above 85%. Meiosis in parental tetraploid strains was characterized by the presence of quadrivalents in addition to a complementary number of bivalents. The average chiasmata frequency per chromosome was 0.59 and pollen fertility was generally about 80%. Tetraploid interspecific hybrids also had quadrivalents, normal meiosis, and high pollen fertility. Close genetic relationships between the diploids and between the tetraploids are indicated, and geographical, ecological, and seasonal barriers to gene exchange exist. Attempts to obtain hybrids between diploids and tetraploids were successful in a few cases. The hybrids were tetraploid and had normal meiosis and fertility similar to parental and F₁ tetraploids. Their origin was by the union of unreduced gametes of the diploid female parent and normal pollen from the tetraploid parent. On the basis of chromosome homology, normal meiosis, plus high fertility exhibited in the diploid, tetraploid, and diploid X tetraploid interspecific hybrids, these species of Grindelia are considered to be a part of an autopolyploid complex. Gene exchange between diploids and diploids, tetraploids and tetraploids, and diploids and tetraploids is possible. Tetraploid G. camporum may have originated by hybridization between G. procera and diploid G. camporum with subsequent doubling of chromosomes and selection for the combined characteristics of the diploids.     

Extensive citrus triploid hybrid production by 2x?×?4x sexual hybridizations and parent-effect on the length of the juvenile phase

The citrus fresh market demands the production of seedless citrus fruits, as seedy fruits are not accepted by consumers. The recovery of triploid plants has proven to be the most promising approach to achieve this goal, since triploids have very low fertility, are generally seedless and do not induce seeds in other cultivars by cross pollination. Triploid plants can be recovered by 2x?×?4x sexual hybridization. In this work, we present an effective methodology to recover triploid plants from 2x?×?4x hybridizations based on in vitro embryo rescue, ploidy level analysis by flow cytometry and genetic origin of triploid plants. The pollen viability of diploid and tetraploid citrus genotypes was analyzed by comparing the pollen germination rate in vitro. The pollen viability of tetraploid (doubled-diploid) genotypes is generally reduced but sufficient for successful pollination. Triploid embryos were identified in normal and undeveloped seeds that did not germinate under greenhouse conditions. The influence of parents and environmental conditions on obtaining triploid plants was analyzed and a strong interaction was noted between the parents and environmental conditions. The parental effect on the length of the juvenile phase was also demonstrated through observations of a large number of progeny over the last 15?years. The juvenile phase length of the triploid hybrids obtained with 'Fortune' mandarin as female parent and tetraploid 'Orlando' tangelo as male parent was shorter than the juvenile phase obtained with a clementine as female parent and tetraploids of 'Nova', 'W. Leaf' and 'Pineapple' male parents. Key message Effective methodology to recover citrus triploid plants from 2x?×?4x sexual hybridizations and the parental effect on the length of the juvenile phase.     

Apoptosis-like degradation of pollen grains in siberian squill related to the lack of low temperatures during their development

Low temperature is necessary for completing the cycle of anther development in early-spring ephemeroids. Pollen grains were investigated in Scilla sibirica L. growing under natural conditions in a park and taken to a greenhouse for the cold season. In control plants grown in the open ground, the development of microspores was normal. The plants transferred to the greenhouse sharply differed from control material. By the middle of November, the aggregations of condensed chromatin were formed near the nucleus envelope of their generative cells. In some mitochondria, the system of cristae was reduced. In December, all the mitochondria showed a reduced system of cristae. In the matrix of some mitochondria with ruptured envelopes, rather large vesicles were detected. In the vegetative and generative cells, autophagic vacuoles were formed. They also occurred in the nucleus itself; as a result, the cells of pollen grains became much more vacuolated. In January, only the products of protoplast lysis were detected in all the pollen grains. It was concluded that the lack of low temperatures induced apoptosis-like degradation of the cells of Siberian squill pollen grains.